L
JOURNAL OF
ENTOMOLOGY AND
ZOOLOGY
VOLUME XII. 1920
PUBLISHED QUARTERLY BY THE
UEPAR'l MENT OF ZOOLOGY OF POMONA COLLEGE
CLAREMONT, CALIFORNL^, U. S. A.
CONTENTS OF VOLUME XII
Volume XII. Number 1
Chamberlain, Ralph V.
Xow Ciililiirnia SpidiTS. 1.
Centipedes and Millepedes from
Near Claremoiit, 24.
Spiders from Clarcim>iit-L:iguiia Ki-
gioii. 25.
H Iton. William A.
Central Nervous System of My-
filus Caliloriiianus, 27.
Chamberlain, Ralph V.
.\ntos fill till- Sipunculicia of La-
Kuna Beach. 30.
Campbell, Arthur S.
Central Nersous System of a Cen-
pcde. 69.
Corwin, Genevieve
Mii.r<iMO|iic Studies of the Water
of the Claremont-LaKuna Re-
gion, 72.
Case, Susie
General Reactions of a Centi-
pede, 79.
Hilton. William A.
Notes on the Central Nervous
System of a Kree-Liviin.i; Ma-
rine Nematode. 82.
Volume XII. Number 2
Munz, Philip A.
A Study of the I'ood llal.il^ i.f
the Ithacan Species of .\nur;i
During Transformation. 33.
Hilton. William A.
The Central Nervmi- S\vtiiii ni
Three MivaKis, .v
Volume XII. Number 3
Cowles, Raymond B.
.\ List and Some Notes on th'-
I.izarcls and Snakes Represented
in the Pomona College Museum.
()3.
)IUton. William A.
The Central Nervous System i^f
an Unknown Species of Marine
Leech. 67.
Volume XII. Number 4
Alexander. Charles P.
■Vew Species of Crane-Flies from
the l"nitcd States and Canada,
Hilton. W. A.
Notes on Pacific Coast Pycyno-
Konids, 93.
Caldwell. J.; Durant, W.
I ca NJusica. '*4.
Caldwell. J.
Lepidopia Myops. 9.5.
Lorbeer. Howard
l'>eniita .Xnaloga. %.
Hilton, William A.
The Nervous Svstein and Sen<'-
Organs. 1. II and III. 1 to II
INDEX TO VOLUME XII.
Ak-xaiulcr, C. I'., 85. Leech. 67.
Aiuira, 33. Lepidopia myop.s. 9.S.
Case. Susie, 79. Lizards, 63.
Caldwell, j., 94, 95. Lorbeer, H., 96
'Jampbell, .\. S.. 69. Microscopic life. 72. 74, 7(i.
Centipede, 24, 69. 7". Millipedes, 24.
Central nervous sy.-.teni, 1, 14. 27. 57, Mytilus. 27.
67. 09. 82. Munz. I'. .A., 33.
Clianilicrlain. K. V., 1. .?(!. Nematode, 82.
Cowles, R. B., 63. Nervous system, 57, 67, 68, 82.
Corwin G.. 72. Pycnogonids, 93.
Crane-rties, 85. Spiders, 1, 25.
Durant, W.. 94. Sipunculida, 30.
Kreniita analoga, 96, Snakes, 63.
Food liabits, 33. Toad, 78.
Hilton, W. A., 27. 57. 67, 82. 93. Uca nnisica. 94.
VOLUME TWELVE NUMBER ONE
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
MARCH, 1920
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT 0/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
NEW CALIFORNIA SPIDERS— «a/;)A V. Chamberlain 1
CENTIPEDES AND MILLEPEDES FROM NEAR CLAREMONT 24
SPIDERS FROM CLAREMONT-LAGUNA REGION 25
CENTRAL NERVOUS SYSTEM OF MYTILUS CALIFORNIANUS
William A. Hilton 27
NOTES ON THE SIPUNCULIDA OF LAGUNA BEACH
Ralph V. Cha mherlain 30
Entered Claremont, CaK.Post-Office Oct. 1, 1910. as second-class matter, under Act of Congress of
March 8, 1878
Journal of Entomology and Zoology
EDITED BY POMONA COLLEGE, DEPABTMENT OF ZOOLOOY
Subscription $1.00 to domestic, $1.25 to foreign countries.
This journal is especially offered in exchange for zoological
and entomological journals, proceedings, transactions, reports
of soL'ieties, museums, laboratories and expeditions.
The pages of the journal are especially open to western ento-
mologists and zoologists. Notes and papers relating to western
and Californian forms and conditions are particularly desired,
but short morphological, systematic or economic studies from
any locality will be considered for publication.
Manuscripts submitted should be typewritten on one side of
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figures, etc., should be written on separate sheets. Foot notes
and figures should be numbered consecutively throughout. The
desired position of foot notes and figures should be clearly
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Figures should be drawn so that they may be reproduced as
line cuts so far as possible. An unusually large number of half
tones must be paid for in part by the author. Other more
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Authors of articles longer than a thousand words will receive
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Manuscripts should be sent by express or registered mail.
Address all communications to
The Journal of Entomology and Zoology
William A. Hilton, Editor
Claremont, California, U. S. A.
cJ— ^A.<l_jt O^-^
New Californian Spiders
RALPH V. CHAMBERI.IN
The new spiders described below were found recently while identifying a col-
lection from Claremont received from Prof. Hilton and one made by the writer in
the same region in 1909 and 1913. A few forms from otiier localities noted in mak-
ing comparisons are also included.
AVICULARllDAE
llexiira fut-va sp. nov.
Carapace and sternum with labium and endites yellow of light reddish cast, un-
marked excepting for the solid black interocular area. Legs pale yellowish brown
without the reddish tinge. Chelicerae typically a little darker than the carapace.
Abdomen grey above and either wholly unmarked or sometimes showing a short
median longitudinal pale line at base; venter paler excepting toward the spinnerets,
where darkened; spinnerets pale brown like the legs. Chelicerae long, clothed
above on mesal portion with long setfe which are more abundant on the anterior face
below, .■\nterior lateral eyes much the largest, less than their long diameter apart,
scarcely three times the diameter of the medians. Anterior median eyes about their
radius apart, between two-thirds and three-fourths the diameter of the posterior
medians, which are smaller than the posterior laterals. Tibife I and II armed beneath
with 3-1-1 spines, the two unserried spines being at distal end. Metatarsi I and II
armed beneath with 3-3 spines. Spinnerets with articles proportioned much as in
p'ura, the tcrmiiial article being pointed and subaimulate, but the length rather shorter
than the width of tite abdomen and much shorter than its length.
Length, 9 mm. Length of ccphalothorax, 4 mm. Length of tib. -)- pat. I, 3 mm.;
of tib. -|- pat. I\', 3.2 mm.
Type— M. C. Z. 380. Claremont.
A much lighter colored species than />iirii, the genotype, and differing in the
much shorter spinnerets, in having 3-3 spines instead of 2-2 below on metatarusus I,
in having the anterior lateral eyes scarcely three times instead of more than four
times the diameter of an anterior median, in the proportionately broader endites, etc.
Nemesoides gen. nov.
Pars cephalica of moderate size. Fovea thoracica moderate, recurved. Anterior
row of eyes procurved, median eyes much smaller than the laterals. Laterel eyes
on each side less than their radius apart, the anterior scarcely larger than the pos-
terior. Rastellum of chelicerE well developed, the teeth long and stout. The labium
broader than long, unspined. Endites armed at base with a patch of slender spines.
Sternum with a pair of large impressions united at middle and in transverse line
with them, near, but separated from, each lateral margin a much smaller impression.
Tarsal claws with teeth numerous, in two sinuous series. Tarsi and, in part, meta-
tarsi of first two pairs of legs scopulate. Tarsi of last two pairs of legs spined (male,
genotype.) Metatarsus IV shorter than tibia IV. Superior spinnerets large, four-
jointed, the distal joint short, rounded, shorter than the third and much shorter than
the second. Tibia I of male with spur.
Genotype — A', hespera sp. nov.
This genus falls in Simon's group Nemesicse in its more restricted sense.
2 Journal of Entomolog>' and Zoologj'
Semrsoides hcspera sp. nov.
Male — Carapace, sternum, labium and endites and legs yellowish. C'helicerae
darkened distad bv the black teeih of the rastellum. Abdomen yellowish beneath;
light brown above, with three longitudinal rows of short, black, transverse marks.
Chelicerr long and rather slender, extending almost directly forward, not at all
geniculate, the lower teeth of rastellum stout. Anterior row of eyes procurvcd in
such manner that the line tangent to the lower edges of the median eyes passes
through or near the centers of the laterals; lateral eyes with diameter twice that
of the medians; median eyes their diameter apart. Anterior and posterior lateral
eyes equal or very nearly so, separated by less than lialf their radius. Posterior
median eyes nearly of same size as the anterior medians from which separated by
their radius, closer to the posterior laterals. Tibia I in male with spur or process;
strongly spined ; a series of long, stout spines along each side, fewer smaller ones
beneath, typically a short, oblii|ue row of four close-set and especially stout spines
at the ectoventral corner of the distal end. Metatarsus I with a strong angle, or
process, at middle of the ventral edge. Palpal organ as shown in pi. 1, fig. 1.
Length, 10 mm. Length of cephalothorax, 5 mm. Length of tib. + pat., 1.5 mm.;
of tib. + pat. IV, 5.5 mm.
Type M. C. Z. 379. Clarcmont. \Vm. A. Hilton coll.
niCTVNID.lC
I miiiiKihius nigrellus sp. nov.
Fftnalr — Carapace dusky chestnut to nearly black. Sternum solid black. Labium
and endites black or blackish excepting across tips. Legs dusky brown, the femora
darker, blackish. Abdomen above and laterally blackish brown, the background black
lightened by numerous minute yellowish dots; venter mesally immaculate black.
Anterior median eyes their diameter apart, once and a half as far from the laterals.
Posterior row of eyes but little longer than the anterior; median eyes nearly twice
their diameter apart, and almost two and a half times their diameter from the
laterals. Area of median eyes wider behind than in front and longer than wide. Tibia
I unarmed. Anterior metatarsi well spined beneath. Tibia IV with four spines
beneath, these in a longitudinal line with an extra one at distal end. Tibir III and
IV with a small spine at the base above, in this differing from the other known
North American species. Lower margin of furrow of chelicerx armed with two
teeth. Epigynum a plate subcordate in outline with a median longitudinal band ex-
tended laterad on each side behind.
Length 6 mm. Length of tib. -f pat. I, 3 mm.; of tib. - pat. IV, the same.
Type— M. C. Z., 374. Cal.. Claremont. Prof. \Vm. A. Hilton.
Pnrauximui gen. nov.
Resembles .Auximus in eye characters, but eyes of both rows nearlv e(|uidistant.
It differs in having the lower margin of the furrow of the chelicerr armed with
eight teeth, instead of four or five, of which the most distal instead of the most
proximal is largest; upper furrow with three teeth of which the median is largest.
.\ notable feature of the genus is that the patella of the male palpus, at least in the
genotype, bears a stout apophysis.
Crnnlyff — /'. tar.lnliii sp. nov.
Pomona College, Claremont, California 3
Parauximus tardatus sp. nov.
Male — Carapace dusky over light brown. Legs with somewhat obscure dusky
annuli over yellow. Labium and endites chestnut, pale across tips. Chelicerr dusky
chestnut. .Abdomen dark over sides, dorsally a pointed mark outlined in black from
base to middle, followed by a series of mesally connected chevron marks. \'enter
immaculate light grey with an angular extension from the dark of each side just in
front of the spinnerets, the two processes not meeting in the middle line. Lower
margin of the furrow of the chelicers bearing four large teeth and proximad of
these four smaller ones. Anterior median eyes very small, rather less than half the
diameter of the laterals, near their diameter apart and about the same distance from
the laterals. Posterior row of eyes straight; median eyes smaller than the laterals.
About their diameter apart and the same distance or a little less from the laterals.
Anterior laterals larger than posterior laterals and separated from them by about a
radius of the latter. Tibice and metatarsi I and II armed beneath with three pairs
of spines.
Palpus as shown in plate 1, fig. _. Patella with a stout apophysis bearing distally
numerous spines.
Type— M. C. Z. 377. Claremont.
Readily distinguishable by the characters of the eyes and the structure of the
male palpus.
Auximus pallescens sp. no v.
Female — A species in appearance much resembling the preceding, though typi-
cally paler with the carapace and legs much more yellow. Sternum yellow. Labium
chestnut, pale across tip, the endites lighter; also distally pale. Abdomen colored
somewhat similarly to that of the preceding species, but the dorsal markings in the
type indistinct. The species is easily distinguished from the preceding by its much
larger anterior median eyes, which equal or nearly equal the laterals and obviously
exceed the posterior medians and which are separated from each other by rather
less than their radius and from the laterals by not more than once and a half their
diameter. Posterior median eyes separated by near once and two-thirds their diameter
and from the laterals by twice and a half their diameter, the laterals much larger.
Lateral eyes on each side separated by their radius or less. Lower margin of furrow
of chelicera armed with four teeth. TibijE I and II and metatarsi I and II each
armed beneath with three pairs of spines. Epigynum, apparently not quite fully
chitinized, shown in plate 1, fig. 3.
Length 12.5 mm. Length of cephalothorax, 6 mm. Length of tib.-fpat. I, 5.7
mm.; of tib. + pat. IV, the same.
Type— M. C. Z. 376. \Vm. A. Hilton coll.
Auximus latescens sp. nov.
Female — Carapace pale chestnut tending to testaceous in posterior and lateral
regions. Legs testaceous to brown, the anterior ones often of slight chestnut cast.
Sternum pale chestnut and the endites and labium darker chestnut. Chelicera dark
chestnut or mahogany. The abdomen above is dark brown to blackish, with a pos-
teriorly pointed pale mark reaching from base to middle followed by a series of pale
chevron marks and on each side of it with usually three pale spots, which may be
more or less connected with it or sometimes a short light line each side; venter grey-
4 Journal of Kntoinology and Zoology
ish brown to yellowish with two rather wide longliludinal dark stripes which are
but narrowly separated on each side from the dark of the sides. Anterior lateral
eyes with diameter once and two-thirds that of the medians; median eyes about livc-
sixfhs their diameter apart, twice and a half their diameter from the laterals. Lower
margin of furrow of chelicera with four teeth, of which the most proximal is largest.
Tibii I and II armed with five spines, one at base, two sub-median and two apical.
Epigynum as shown in plae 1, fig. 4.
\lalf — Carapace and legs somewhat paler than in the female. Eves less widely
separated. Palpal organs as represented in plate I, fig. 5.
Type— M. C. Z. 372. Cal.: Claremont. Type taken by the author in 1909. Para-
types take in 1913. Also In 1918 coll. of Prof. Hilton.
The genus to which this and the preceding species belong, known from South
.America and the Atlantic Islands, has not previously been recorded from North
.America.
Diityna mians sp. nov.
Female — Pars cephalica yellowish, other parts of carapace brown to fuscous.
Sternum yellowish, sometimes a little dusky, with the labium similar, but endites
ordinarily paler. Legs not annulate in the types though the femora may be slightly
darkened and the tibia and metatarsus show vague darkening at distal end. Ab-
domen above yellowish, with a dark spot in front of middle from which some fine
dark lines railiatc and anastamnse to form a network, the median longitudinal line the
best developed of these; typically three pairs of widely separated dark spots on
posterior portion, but these often broken or indistinct. Venter darker, sometimes a
median yellow spot in front of the cribellum with one in each edge of dark area.
.Anterior row of eyes straight; median eyes their diameter or a little more from the
laterals, farther from each other. Posterior eyes nearly equidistant. Area of median
eyes wider behind than in front. Epigynum, plate 3, fig. S.
Type— M. C. Z. 385.
Cal.: Los Angeles Co. ( R. V. Chamberlin); also northern part of stale (Peck-
ham coll.).
Has resemblance to P. laliarata, occurring in the same localities, but easily dis-
tinguished by the structure of the epigynum and the more widely separated eyes.
scvTomn.K
Plrttmtrys suftrrntins sp. nov.
Frmalf — Differs at sight from /'. aislanfa Simon, which occurs in the same region.
In Its much longer legs, lighter, more dilute chestnut, carapace, and the proportion-
ately shorter and higher abdomen. The legs are brown, of less chestnut cast, with the
first ones not ronirasting by deeper, fuscous color. Sternuin pale chestnut like the
carapace. .Abdomen cinereous of slight greenish cast, with pale median mark on
dorsum at base. The anterior row of eyes is longer than in caslanfa with the
lateral eyes comparatively smaller , their diameter not exceeding once and a half
that of the medians; median eyes atwut their radius apart, much farther removed
from the laterals than in eatlanra, the distance being from two and a half to three
limes their diameter. Posterior row of eyes distinctly a little recurved Instead of
straight, with the median eyes larger than the laterals instead of a little smaller,
Pomona College, Clareniont, California 5
separated by their longer diameter or more, a little nearer to the laterals. The
trapezium of median eyes is much wider in proportion to the length than in castanea.
Tibia 1 with five to seven long, widely separated spines on ventral side, of which none
are paired or, rarely, eight present with Hvo at distal end. Spines under metatarsus
I shorter, very numerous.
Male — Tibia I of palpus withoul apophysis at distal end. Palpus represented in
plate 2, fig. 1.
Length of female, 11 mm. Length of cephalothorax, 5 mm. Length of tibia 4 pat.
I, 6.4 mm. ; of tib. + pat. IV, 4.7 mm.
Type— M. C. Z. 36S. Cal.: Los .Angeles, Claremont. R. \. Chamberlin coll., 19119.
Wni. .\. Hilton coll, 191S.
DR.'KSSID.E
Drassodes teles sp. nov.
FemtiU — Carapace and sternum with entlites and labium testaceous, and legs
yellow. Chelicera; darker brown or pale chestnut. Abdomen ventrally clear yellow
in front of the genital furrow excepting the dark epigynal area; behind the furrow
dusky grey over a yellow background ; dorsally dark olive grey due to dense clothing
of hair. Upper margin of furrow of chelicera with three teeth of which the median
is largest; lower margin with two small teeth. Anterior row of eyes rather stronglv
procurved; median eyes a little more than their diameter apart and a little more
than their radius from the laterals, which are nearly their diameter from lower edge of
clypeus. Posterior row of eyes scarcely procurved, much longer than the anterior
row; lateral eyes smaller than the anterior laterals from which separated by once
and a half the diameter of the latter; median eyes oblique, scarcely more than their
long radius apart, twice their long diameter and nearly three times their lesser diameter
from the smaller laterals. Tibiae I and II armed beneath with but a single spine,
which is attached a little distad of middle and toward the mesal side. Metatarsi I
and II with a single spine beneath, this at base. All tarsi scopulate. Anterior
metatarsi, and metatarsus III at distal end also, scopulate. Epig\ni'm represented in
pi. 2 f. 2.
Length, 10 mm. Length of cephalothorax, 4.5 mm. Length of tib. + pat. I, 4.5
mm.; of tib. -f pat. IV, 5 mm.
Type— M. C. Z. 360. Cal.: Claremont. \Vm. A. Hilton.
An obviously larger species than D. robustus which has a very different epigynum
and bears no spine under tibia I. Only the male of D. californicus is known; but
this may be distinguished from the present species by its difl^erent eye relations; e. g.,
in having the posterior laterals larger than the medians and the latter farther apart.
It also has two pairs of spines under tibia, I which may not be a secondary character.
Scoptop/iaeiis ■vnliititarius sp. nov.
female — Carapace, sternum and legs pale chestnut, the posterior legs and the
coxae beneath more brown and the anterior legs dusky or blackish beyond the femora.
Endites like sternum, the labium and chelicera a darker chestnut. Abdomen blackish
grey above and laterally, with a faintly indicated pale mark at base above; venter
yellow in front of genital furrow and dusky greyish yellow behind it, with a pair of
interrupted longitudinal dark lines. Epigynum blackish. Furrows of chelicerje
unarmed. .Anterior row of eyes procurved; median eyes between one-half and three-
6 Journal of Entomolog\- and Zoology
fourths ilicir diameirr apart, only about one-eighth their diameter from the much
smaller lateral eves and less than their diameter from the lower edge of clypeus.
Posterior row of eyes a little longer than the anterior, a little procurved; median eyes
their diameter or scarcely more apart, closer to the laterals. All tarsi with well
developed scopula: and the anterior metatarsi also scopulate. Tibii I and II each
with a single spine at distal end beneath and metatarsi I and II each with one at
base beneath. For form of epigynum see pi. 2, f. 3.
Length 8.5 mm. Length of cephaloihorax 4 mm. Length of tib. + pat. I, 3 mm.;
of tib. -f pat. IV, 3, 1 mm.
Type— M. C. Z. 361.
Uerpytlus pius sp. nov.
Fftnale — This large form in general appearance resembles //. validus, which is
common in the same region ; but, aside from readily noted differences in eyes and
especially in the epigynum, it may easily be distinguished in having no spines beneath
on tibia I, whereas valiJus has three spines as on tibia II, which is similarly armed in
the present species. Carapace and legs pale chestnut. Sternum and endites similar
but the labium and chelicera; darker. Abdomen grey, densely clothed with hair, as
usual, the type not showing any delinite markings. Hairs of plumose type, as usual.
Posterior row of eyes considerably longer than the anterior, clearly procurved;
median eyes circular, subequal to or scarcely smaller than the laterals, slightly more
than their diameter apart and twice their diameter from the laterals. Anterior median
eyes considerably larger than the laterals, their radius apart, closer to the laterals.
Furrow of chelicera; armed above with three small teeth, below with one. For
epigynum see pi. 2, f. 4.
Length, 11 mm. Length of ccphalothorax, 5 mm. Length of tibia patella I, 4.5
mm.; of tib. -f pat. IV, 5 mm.
Type— M. C. Z. 365.
Cal.: Claremont. R. \. C'hamberlin coll., 1909.
Zelolfs latho sp. nov.
Female — Carapace and sternum reddish yellow, the legs yellow without the red-
dish cast. Endites like sternum, the labium and chelicera: darker. Abdomen grey
without distinct markings. Posterior row of eyes distinctly longer than the anterior,
a little procurved; median eyes elongate, elliptic, very oblique to each other, larger
than the laterals, separated from each other by less than their radius, nearly their
diameter from the laterals. Anterior median eyes smaller than the laterals, about
their radius apart, not more than half as far from the laterals. Lateral eyes on each
side separated by more than their radius but less than their diameter. Tibia I
unarmed beneath, metatarsus I with a ventral spine at base. Tibia II beneath with a
submedian spine, metatarsus II with a spine at base. Form of epigynum represented
in pi. 2, f. 5.
Length, 6.5 mm. Length of ccphalothorax, 2.9 mm. Length of lib. -} pat. I, 1.4
mm.; of tib. -f pat. IV, nearly the same or slightly less.
Type — M. C. Z. 367. Claremont.
'/.rlntes irritnni sp. nov.
Male — Carapace, slcrniun, legs, and mouthpnrts dusky over a yellow background.
Pomona College, Claremont, California 7
the anterior tibije more blackish than the posterior. Abdomen greyish black. Posterior
row of eyes but little longer than the anterior, slightly procurved; median eyes
broadly slightly obovate, much larger than the laterals, separated from each other by
less than their radius, twice as far from the laterals. Anterior median eyes very
much smaller than the laterals, to which they are very close, separated from each
other by their diameter. Tibia I armed beneath with a single submedian spine;
tibia II armed beneath with three spines, two of these being submedian and at slightly
ditfereiit levels and one sub-basal. Palpus as shown in pi. 2, f. 6.
Length, 5.1 mm. Length of cephalothorax, 2.25 mm. Length of tib. + pat. I, 2.1
mm.; of tib. -|- pat. I\', 2.5 mm.
Type M. C. Z. 366. Claremont.
Zelotes gynelhiis sp. nov.
Female — A dark colored species having the general appearance of Z. niger but
readily distinguishable in its smaller and very differently formed epigynum, etc., and
from other species also by that character and those of the eyes. Carapace black of
slight chestnut cast, shining. Legs dusky mahogany or the proximal joints, especially
of the anterior pairs, solid black. Sternum dusky chestnut, the labium and endites
similar. Abdomen greyish black above, paler beneath, without markings. Posterior
row of eyes very slightly procurved, considerably longer than the anterior row;
median eyes nearly their diameter from the laterals and a little nearer to each other.
The anterior median eyes are characteristically very small, being greatly exceeded
by the laterals from which separated by not more than half their radius, separated
from each other by once and a half or more their diameter. No ventral spines on
tibiae I and II or on corresponding metatarsi. For form of epigynum see pi. 3, f. 1.
Length, 8 mm. Length of cephalothorax, 3.1 mm. Length of tib. + pat. I, 2.9
mm.; of tib. -f pat. IV, 3.4 mm.
Type— M. C. Z. 363. Cal.: Claremont.
Zetott's t'thops sp. nov.
Male — Carapace and legs brownish yellow, the sternum clearer yellow. Labium
darker than sternum, the endites like sternum. Chelicerae brown. Abdomen grey.
The species seems readily distinguishable from those described previously from North
America in the atypical character of the eyes and endites. The posterior row of
eyes, which is straight, not at all longer than the anterior, the eyes all being close
together, the medians but slightly separated and but little farther from the somewhat
smaller laterals. The anterior row of eyes procurved with the laterals but little more
than their radius removed from the edge of the clypeus ; the median eves, which are
much smaller than the laterals, separated by but little more than their radius and much
closer to the laterals. Lateral eyes on each side much nearer to each other than the
medians, separated by less than their diameter. Chelicerse armed above with three
small teeth, below with two. The endites are characterized by having the palpus
inserted at or a little distad of the middle, obviously farther distad than usual. Tibia
I and metatarsus I unarmed beneath; tibia II also unarmed beneath but metatarsus II
with two spines in longitudinal line beneath. .Anterior spinnerets large, much exceeding
the posterior.
Length of not fully mature male type, 6 mm. Length of cephalothorax, 3.1 mm.
Length of tib. + pat. I\', 3.4 mm.
Type M. C. Z. 362. Cal.: Claremont.
8 Journal of Entomology and Zoolog>
PHOLCIP.l
Psilof/iorus californi,r sp. nov.
Carapace, slcrnum. and legs yellow or the carapace and legs proximally of pale
brown casi ; the femora proximally and the patella and tibis at ends often tinged
with bright red. The head and the furrows commonly darker than other parts of
carapace, with the eyes enclosed in black. The abdomen to the naked eye appears grey,
commonly of a greenish tinge; under the lens it shows on the sides numerous light,
somewhat silvery, spots and above a basal pale mark, with several pairs of dark spots
enclosed by the light ones and often more or less subdivided. Posterior row of eyes
straight; the median eyes nearly their diameter apart, their radius or a little more
from the anterior lateral eyes, and three-fourths their diameter from the anterior
medians. Anterior eves in a strongly procurved row, with the medians much the
smaller, as usual. In the male the apophysis on the chelicera is attached near the
middle of the anterior face and projects directly downward or a little forward of
downward; it is smaller than in corniiliu and differs also in position and form from
that in /<ullulu3. (PI. 3, f. 2.) The species is most readily recognized by the structure
of the male palpus, which is represented in pi. 3, f. 3.
Length (male), 3.2 mm. Length of femur I, 4.S mm.; of femur IV, 3.S mm.; of
tib. -f pat. I, 5 mm.; of tib. -f- pat. IV, 4 mm.
Type— M. C. Z. 370.
Cnl ('lnr<-m..„i R V Chamberlin coll., 19(19. .Also \Vm. .A. Milton oill.. 1918.
TIIERIOIin.T,
l.illiyfilianirs mimiiiJrs sp. nov.
h'emale — Carapace reddish brown or chestnut, darker on lower part of sides and
svilh an obscure median longitudinal dorsal line on pars cephalica at least. Sternum
chestnut, sometimes nearly black. Legs chestnut, with anterior tibiae darker. Cheli-
cer.T, labium and enditf> darker, almost mahogany, .'\bdomen in general silvery white,
with n cllr^e network of tine brown lines; dorsum typically with four pairs of dark spots
of which the most caudal arc united; a narrow, brown hastate mark along middle, a
brown stripe on anterior face and extending caudad along each side where it bifurcates,
a series of obli(|ue lines uniting the two branches in the caudal region; venter covered
»vith a network of dark lines and spots. Anterior row of eyes nearly straight or
slightly procurved. Anterior median eyes smaller than the laterals, their diameter or
more apart and slightly farther from the laterals. Lateral eyes on each side narrowly
separated, obviously closer to each other than in lornllalus. equal. Posterior row of
eyes slightly procurved. Posterior median eyes their diameter apart, nearly twice as
far from the ei|ual laterals. The species is easily separable from /.. corollalus, which
it superficially resembles, by the strongly different form of the epigynum as well as by
the difference in eye arrangement noted above. See pi. 3, f. 4.
Length, 7.5 mm. Length of cephalothorax, 2.9 mm. Length of tib. -f pat. 1, 3.4
mm.; of tib. -|- pal. IV, 3.2 mm.
Type— M. C. Z. 340. Oregon: Portland. S. Henshaw coll., June 19, 1882.
ARCilOPin.K
.Iraiifti gnsnijiina sp. nov.
Ffinalf — This species falls In the group with longitudinal thoracic furrow, the
Pomona College, Claremont, California 9
anterior femora armed beneath with a double series of numerous stout spines, and the
abdomen broadly triangular-oval in outline {Neoscona in part.) In coloration it
differs from .-I. utahana Chamb., e. K-. in having the anterior tibia: and metatarsi only
biannulate instead of triannulate, the median annulus being absent, while the femora
have an annulus only at the distal end. In the type the carapace is somewhat dark-
ened in a median longitudinal stripe and may have been blackish in life. Thorax
blackish at sides. Abdomen in general light yellowish; on posterior portion above a
black line with posterior end bifurcating, and a black line on each side also running
caudad from anterior end of the median line; on sides a series of brownish, parallel,
subvertical lines; venter not unusually black as it is in ulaliana. The scape of the
epigynum instead of curving evenly with convexity ventrad, is straight to the distal
end which is bent abruptly ventrad instead of curving dorsad as in vertehrata. This
bending may in part be an artifact as the abdomen in the type was shrunken firmly
against the end of the scape. See pi. 6, f. 6.
Length, 14 mm. Length of abdomen, 11.5 mm.; width, 9.6 mm. Length of
cephalothorax, 6.6 mm. Length of tib. -f- pat. L "■- min.; of tib. -}- pat. IV, 6.5 mm.
Type— M. C. Z. 388. Cal.: Desert region.
THOMISID,^
Thanalus retentus sp. nov.
Female — Carapace with a chocolate colored band on each side above a pale
marginal stripe, with a broad median dorsal pale stripe embracing typically a darker
median longitudinal mark which bifurcates at the posterior border of head and is
continued forward as interrupted dark lines, a median dark line also present betweeen
these branches. Lower median region of clypeus pale. Sternum yellow, densely dotted
over borders, or sometimes over entire surface, with minute dark spots. Legs brown,
lined and mottled with black, the joints showing some clearer longitudinal lines par-
ticularly on the femora. Abdomen above yellowish with a dark colored basal sagittate
mark reaching to middle or indistinctly continued bej'ond in an interrupted median
line; on posterior region a dark area showing several chevron marks united on each
side in a line or band with wavy exterior edge; typically the venter shows two
narrowly separated median black lines united in an acute angle in front of spinnerets
and ectad of this on each side another dark line. Posterior row of eyes strongly
recurved, as usual, the median eyes scarcely nearer to each other than to the laterals
(cir. 14:15). Area of median eyes narrower in front than behind, longer than wide
in about ratio 2il:17. Anterior medians twice as far from each other as from the
laterals. Epigynum as shown in pi. 6, f. 5.
Type— M. C. Z. 389.
Claremont. A common species in this region.
This form is readily distinguishable from eolor/iih-niis, with which it has hereto-
fore been confused, by the obviously different form of the epigynum.
AGELENTD.?^
Agelena rua sp. nov.
Male — Carapace with the sides dark, as usual, the median band yellow. Sternum
dusky over yellow with a clear median longitudinal line. Legs light yellow, obscurely
10 Journal of Entomolog>' and Zoolog\'
annulate witli dark. Cheliccra; pale brown. Dorsum of abdomen dark grey along
sides, the median region light reddish ivilh a series of yellow spots along each edge;
sides of abdomen yellowish grey lightly spotted with black; venter limited on each
side by a longitudinal dark line, the intervening region almost immaculate. Posterior
eyes equidistant, not fully their diameter apart. Anterior median eyes much smaller
than the laterals, near their radius apart, a little nearer to the laterals. Palpal organ
represented in pi. 4, f. 1.
Length, 7 mm. Length of cephalothorax, 3.2 mm. Length of tib. -j- pat. I, 4.5
mm.; of tib. -f pa'- '^'i +•" nun-
Type— M. C. Z. 384. California: Catalina Id.: Avalon Bay. \Vm. A. Hilton
coll.. Aug. 25, 191S.
Distinct from other North American species especially in the structure of the male
palpus.
CLIBIONID/E
Ol'ioj schistus sp. nov.
A species approaching O. peiiinsulaniis, known from Lower California, but dif-
fering in coloration and various details of structure. While in peniniulantis the
carapace, labium, endites, chelicerc and legs are uniformly immaculate pale yellow,
in the present species the legs are darkened by numerous minute, dark, somewhat
purplish, spots which show a tendency to condense into an irregularly defined annulus
at proximal end of tibii; similar but fewer dots occur on carapace and chelicer.T,
but the sternum is immaculate. .Abdomen also very obviously darker and differently
marked, being densely spotted and streaked on the sides with blackish and less strongly
so above and below, the dorsum with a clear sagittate mark at base, followed by a
series of short chevron marks united along middle by a black line which is furcate
at its anterior end. Anterior eyes obviously larger than the posteriors; anterior median
eyes their diameter from the laterals and a little farther from each other, the eves
being more widely separated than in pcninsulanus. Posterior rows of eyes a little pro-
curved instead of straight, and the eyes much more widely separated than in the
species mentioned, the medians being three times their diameter apart and as far or
nearly as far from the laterals. Epigynum decidedly larger proportionately, with the
outer ridges posteriorly more thickened and elevated with reference to the inner rims,
etc. See pi. 4, f. 2. The palpal organ of male of similar structure but obviously
heavier; the proximal apophysis of tibia larger, distally clavately expanded and trun-
cate instead of being distally pointed with the setose edge long and oblique; the
anterior apophysis also differing as shown in pi. 4, f. 3.
Length of female, 10.5 mm. Length of cephalothorax, 4.S mm. Length of lib. -f
pat. 1, 6.8 mm.; of tib. -f pat. IV, 6 mm. A male with cephalothorax 4.8 mm. long
has tib. -f pat. I, 8 mm. and tib. -f pat. IV, 6 mm. long.
Type— M. C. Z. 3 54.
Cal.: Clarcmont. R. V. Chamberlin coll. .\\>» Win. .\. Hilton 1918 coll.
.Inyp/inrna irfhriipina sp. nov.
jWrt/c— Carapace and legs dull yellow, a dusky band along upper part of each side
of the former. Sternum, labium and endites also yellow, the chelicerx brown. Abdo-
men dull grey of slight yellow cast; dorsum with a few dark spots, the sides with
more numerous dark spots anil streaks; venter with some spots on posterior portion.
Pomona College, Claremont, California 1 1
dusky in front of genital furrow. Armature of chelicerae normal. Anterior row of
eyes straight; eyes less than their diameter from lower margin of clypeus. Anterior
median eyes obviously smaller than the laterals, rather less than their radius apart,
closer to the laterals. The lateral eyes on each side their radius apart. Tibis I and
II armed beneath with three pairs of long spines, the corresponding metatarsi with
two pairs. Coxae of third and fourth and femora of third legs densely spinulose
beneath. Furrow of posterior spiracles a little behind middle of abdomen. Palpus pi.
4, f. 4.
Length, 5 mm. Length of cephalothorar, 2.5 mm. Length of tib. -f pat-, 2.6 mm.;
of tib. + pat. IV, the same or nearly so.
Type — M. C. Z. 353. Cal.: Claremont. Pomona College coll.
Anyphitna mens sp. nov.
Male — Carapace and legs yellowish, the legs with some obscure dusky markings.
Sternum, labium and endites yellow. Abdomen yellowish grey; immaculate beneath;
streaked and spotted with brown over the sides and the lateral portion of the dorsum;
dorsum posteriorly with two or three rows of spots more or less confluent into chevrons,
preceded by a pair of spots, the anterior median region of dorsum immaculate. Arma-
ture of chelicera typical. Clypeus not quite as wide as diameter of anterior eyes.
Anterior row of eyes straight. Anterior median eyes a little smaller than the laterals,
their radius apart, much closer to the laterals. Posterior eyes equal, obviously longer
than the anterior ones, the row very slightly procurved. Posterior median eyes their
diameter or slightly farther apart. The eyes in general closer together than in incursa,
those of which they somewhat suggest. Tibis I and II armed beneath with two pairs
of spines — one pair basal and one submedian — and metatarsi I and II similarly armed,
the spines in length from about once and a half to twice the diameter of the joint.
Furrow of posterior spiracle rather behind middle of abdomen. Palpus as shown in
pi. S, f. 1.
Type— M. C. Z. 352. Cal.: Claremont. R. V. Chamberlin coll.
Anypluena zina sp. nov.
Female — Carapace yellow, somewhat darker on the sides, as usual. Legs yellow,
marked with a few much interrupted and often obscure annuli, the femora beneath with
a longitudinal row of black dots. Sternum, labium and endites yellow. ChelicerfE
brown. Abdomen yellowish grey; minutely spotted with dark above and over the
sides; venter mostly nearly free from spots, but with a dark line from epigynum to
furrow of posterior spiracle. Clypeus about as wide as an anterior median eye.
Anterior row of eyes a little recurved, .'\nterior median eyes much smaller than the
laterals, not more than their radius apart and much closer to the laterals. Posterior
median eyes and anterior laterals about equal in size, the posterior laterals larger.
Posterior ro%v of eyes slightly procurved. Posterior median eyes a little more than
their diameter apart, a little nearer to the laterals. Lateral eyes on each side more
than their radius but obviously less than their diameter apart. Tibias I and II armed
beneath with three pairs of long spines, none of which are apical. Metatarsi I and II
with two pairs of spines beneath. Furrow of posterior spiracle behind middle of abdo-
men. Epigynum as shown in pi. 4, f. 5.
Length, 6.5 mm. Length of cephalothorax, 2.5 mm. Length of lib. + pat. I, 2.6
mm. ; of tib. + pat. IV, 2.7 mm.
Type— M C. Z. 35L Cal.: Claremont. Wm. A. Hilton coll.
12 Journal of Entomolojj) and Zoology
.Inyph.rna incur sa sp. nov.
>>«!«/»•— Carapace dull yellow, darkened over the sides. Sternum, legs, endites
and labium yellow. Ctielicen chestnut. Abdomen in general yellowish grey, with
a dark stripe along each dorsolateral surface, the two stripes uniting at the spinnerets.
Lower margin of furrow of chelicera bearing the usual series of seven or eight small
teeth. Anterior row of eyes slightly recurved, the eyes not fully their diameter from
the edge of the clypeus. Anterior median eyes only slightly smaller than the laterals,
their radius or scarcely more apart and not more than half as far from the laterals.
Lateral eyes on each side their radius or more apart. Posterior row of eyes procurved,
longer than the first row by about twice the diameter of an eye; eyes subequal to
each other and to the anterior laterals. Posterior median eyes nearly once and a
half their diameter apart and about their diameter from the laterals. Tibia I armed
beneath with two pairs of long slender spines, one pair being basal and one median.
Metatarsus with one pair of spines beneath, these basal. Tibia II armed beneath
with two unpaired spines corresponding to the posterior members of the pairs present
on I. Metatarsus 11 with a pair of spines at base beneath. Posterior spiracle in
front of middle of abdomen. Epigynum as shown in pi. 5, f. -.
Length, 6.6 mm. Length of cephalothorax, 2.8 mm. Length of tib. -f- pat. I, 1.2
mm.; of tib. -f- pat- 'V, 2.9 mm.
Type— M. C. Z. 35n. Claremont. Pomona College Coll.
.Inyplnrna munjclla sp. nov.
Femiilr — Carapace yellow of pale brownish cast, a little darkened <in the sides.
Sternum yellow. Legs of same color as carapace. Abdomen above grey marked with
numerous distinct dark dots, which show a tendency to be arranged in transverse
series; venter paler, almost immaculate, reddish in front of genital furrow, the
epipynum dark. Lower margin of furrow of chelicari armed with a series of seven
or eight small teeth which decrease in si/e proximad. Anterior row of eyes straight,
each removed by more than its radius but less than its diameter from lower tnargin
of clypeus. Anterior median eyes only very slightly smaller than the laterals, their
radius or but little more apart but only slightly separated from the lateral on each
side. Lateral eyes on each side about their radius apart. Posterior row procurved;
eyes very nearly c<|ual in size to the anterior laterals, subequal to each other or the
medians scarcely smaller. Posterior median eyes once and a half their diameter apart,
very nearly their diameter from the laterals. Tibia- I and II and also metatarsi I and
II each armed beneath with two pairs of long apincs. Furrow of posterior spiracle at
middle or slightly behind middle of abdomen. Kpigyninn as shown in pi. 5, f. 3.
Male — Palpal organ as shown in pi. 5 f. 4.
Length of female, 6.8 mm. Length of cephalothorax, 2.9 mm. Length of tib. -f
pat. 1, 4 mm.; of tib. ^ pat. IV, 3 mm.
Type— M. C. Z. 34.S.
Cal.; Claremont. R. V. Chamberlin coll. .Also Wm. .\. Milton cull., 191S.
tnaclirmmis gen. nov.
Cephalothorax similar in form to that of Chemmis. Anterior row of eyes nearly
•iraighl or a little recurved. Anterior median eyes smaller than the laterals. Posterior
row of eves much longer ili.nti ilic anterior row with eves l.irgrr, the medians notably
Pomona College. Claremont, California 13
exceeding the anterior medians, the row typically a little recurved. Area of median
eyes narrower in front than behind where the width almost equals the length. Clypeus
narrow, not exceeding the anterior eyes. Later eyes contiguous. Labium, endites and
sternum essentially as in Chemmis, but the lower margin of the furrow of chelicerae
armed with three teeth instead of two. The anterior tibiae bear similarly five pairs
of long spines beneath ; but the metatarsi bear three pairs instead of two. The
posterior tibiae bear two median dorsal spines instead of wholly laclcing these as in
Chemmis.
Genotype. — A. sober sp. nov.
This genus appears also to include Chemmis unicnlor of Banks from Arizona in
addition to the two species here described. These species are more uniformly colored
than the species of Chemmis; and, in having all markings nearly obliterated, contrast
conspicuously in general appearance with the members of that genus.
Anachemmis sober sp. nov.
Female — Carapace dusky over a brown to light chestnut ground. Legs light brown
to light chestnut, without markings. Sternum light brown or testaceous. Labium and
endites darker, pale across distal ends. Chelicers chestnut. Abdomen dusky brown or
blackish, with a median dorsal light line at base extending to near middle, a light
spot on each side opposite each end and two or three pairs of light spots farther caudad,
but these light marks commonly vague; venter paler, showing a yellowish back-
ground darkened by dark psots. Anterior row of eyes nearly straight; the median
eyes much smaller than the laterals and especially than the posterior medians, their
diameter apart and about half as far from the laterals. Posterior row of eyes a
little recurved, eves subequal; posterior median eyes about their diameter or a little
more from the laterals, three-fourths or less theit diameter apart. Epigynum as shown
in pi. 5, f. 5.
Length up to 10.5 mm. Length of cephalothnrax, 4.5 mm. Length of tib. + pat.
I, 5 mm. ; of tib. -|- pat. IV, the same.
Cal.: Claremont. R. V. Chamberlin. Paratypes from same region also in col-
lection received from Prof. Hilton.
Anachemmis Ji>lichopus sp. nov.
Female — Contrasts in general appearance with the preceding species in its very
long legs and much lighter color. The carapace, sternum and legs are yellowish
brown without markings, but the legs are somewhat darkened over tibias and distal
joints. The abdomen is uniform grey throughout, with no definite markings. The
anterior row of eyes slightly recurved; median eyes much smaller than the laterals,
about their diameter apart, closer to the laterals. Posterior row of eyes straight;
medians smaller than the laterals, about their diameter apart, nearly half as far again
from the laterals. Legs very long. Readily distinguishable by the form of the
epigynum as shown in pi. 5, f. 6.
Male — Palpus shown in pi. 5, f. 7.
Length of female, 10 mm. Length of cephalothorax, 4.5 inm. Length of leg I,
exclusive of coxje, 17 mm.; of tib. + pat. I, 6.5 mm.; of tib. — pat. IV, the same.
Type— M. C. Z. 344. Cal.: Claremont. Win. .A. Hilton coll.
14 Journal of Entomology and Zoology
Samofisilus gen. iiov.
Ccphalochorax wiili general form much as in Trachelas. Sternum broailly trun-
cate anteriorly, pointed at caudal end, margltied. Endites not excavated exteriorly, as
hroad at middle as at distal end; the disloeclal corner rounded. Labium distally
truncate or a little incurved. Lateral eyes on each side well separated, though much
closer together than the anterior and posterior medians. Anterior row of eyes straight.
Anterior medians smaller than the laterals. Posterior row of eyes slightly recurved,
eyes equidistant or nearly so and nearly equal in size, with the laterals equal to the
posterior laterals. Quadrangle of median eyes wider behind than in front. Clypeus
much wider than the anterior eyes. I'pper margin of furrow of chelicera with three
large teeth, of which the median is longest; lower margin with a series of seven or
eight teeth, of which the most proximal ones become reduced in si/e. None of the legs
scopulate and all lacking terminal tenent hairs. Anterior tibia; armed beneath with
four pairs of long spines, the metatarsi with three pairs. Posterior tibia? in middorsal
line with a basal and a subapical spine, and each patella with a median spine at
distal end above, these dorsal spines smaller than the laterals and ventrals.
Gfnolypr. — A', pletus sp. nov.
Samofisilus />lrliis sp. nov.
Ffmalf — Carapace chestnut colored, dusky over the sides, eye region, along striiB
and over clypeus. Legs light chestnut-brown; femora marked with two wide dark
annuli, one at distal end and one submedian, these more or less interrupted above;
patella with annuliis about distal half also interrupted above; tibiae with two broad
annuli, one at distal end and one between middle and base, these sometimes almost
confluent; entire metninrsi dusky or obscurely biannulate. Sternum light chestnut, the
cox.T of legs lighter brown. Chelicera." dusky chsctnut. Labium and endites pale across
tips, elsewher edark chestnut. Sides of abdomen deep brown or blackish, the dorsum
with a series of dark chevron marks ending in the dark of the sides and connected
along the middorsal line, the spaces between them on each side yellowish; venter
grey. Clypeus twice as high as the diameter of an anterior lateral eye. Anterior
median eyes much smaller than the laterals, about their diameter from each lateral eye
and considerably farther from each other. Lateral eyes on each side about their radius
apart. Posterior row of eyes a little recurved. Posterior median eyes scarcely smaller
than the laterals, nearly once and a half their diameter apart and an equal distance
from the laterals. F.pigynum as shown in pi. 6 f. \. The spcrmalheca-, which ordi-
narily show through the integument as black bodies, are not represented in the figure.
Length, 6.5 mm. Length of cephaloihorax, 4 mm. Length of tib. -| pat. I, 4 mm.;
of tib. -f pal. IV, 3.7 mm.
Type— M. C. Z. 346. Cal. : Claremonl ( R. V. Chamberlin coll.; also Pomona
College coll.).
i.vcosin.K
l.ycnsa ferrinilosa sp. nov.
Carapace brown, paler in a supramarginal line on each side, below which the
marginal dark band is interrupted, and in a median longitudinal stripe which narrows
forward and projects in a point between the eyes and again expands between the first
and secoiul rows; a curved line each side of the median stripe just caudad of the eyes
Pomona College, Claremont, California 15
Legs testaceous, without markings excepting tibia IV, which is banded at each end
with blacli, and metatarsus IV, which is darkened at the extreme tip. Sternum, coxeb
and abdomen beneath solid black. Labium and endites black, pale across tip. Cheli-
cerae brown to bright chestnut. Abdomen above testaceous to yellow with a dark
spear-shaped outline over basal part and ending on a chevron mark behind middle,
this followed by a few other chevrons; a number of oblique lines extending out from
basal mark on each side. A black band across each anterolateral corner and extending
along the side where it breaks into streaks and spot; light areas of abdomen clothed
with yellow hair. Upper margin of furrow of chelicera; armed with three teeth; the
lower margin also with three teeth with are stout and subequal. Anterior row of
eyes much shorter than the second, distintly procurved, median eyes their radius or
slightly less apart, an equal distance from the lateral eyes which are decidedly smaller.
Lateral eyes scarcely their diameter from lower margin of clypeus, an equal distance
from eyes of second row. Eyes of second row less than their diameter apart. Quad-
rangle of posterior eyes comparatively long, the cephalothorax being less than three
and a half times as long.
Spines beneath tibiae long and distally very fine. Epigynum small, of form shown
in pi. 6, f. 2.
Length, 16.5 mm. Length of cephalotliorax, 8 mm. Length of tib. + pat. I, 6.6
mm.; of tib. + pat. IV, 7.5 mm. A male with cephalotliorax 8 mm. long has tib. -f-
pat. I, 8 mm. and tib. + pat. IV, 8.5 mm. long.
Cal.; Claremont. R. V. Chambrelin coll.
This species suggests L. concolor Banks of Lower California. It is a smaller
species distinguishable in having tib. -|- pat. IV shorter than the cephalothorax instead
of clearly longer; in having a black band at both ends of tib. IV instead of only at
one end; in not having the femora, metatarsi and tarsi black beneath, etc.
ParJosn tunha sp. nov.
Female — In the types the body is dark throughout, in life clothed with grey hair;
the median dorsal stripe of carapace obscure. Legs black excepting tarsi and meta-
tarsi, which are dull brown, the latter with three black annuli; sometimes the proximal
joints also show the paler color in spots and streaks or in part may be somewhat
annulate. Sternum solid black. Abdomen with integument black above excepting an
obscure pale mark at base; also black laterally, but the venter paler though with a
deep black band between epigynum and spinnerets ; venter in life clothed densely with
grey hairs, the dorsum with grey and reddish intermixed with some black. Anterior
row of eyes slightly procurved, much shorter than the second row ; median eyes their
diameter apart, not fully half as far from the four-fifths as large laterals; the latter
twice their diameter from the edge of the clypeus and decidedly more than their
diameter from the eyes of second row. Eyes of second row fully their diameter, or
slightly more, apart. Two first pairs of spines of anterior tibiae and metatarsi long,
slender, overlapping as usual. Armature of chelicerae typical. Epigynum of the
sternalis type, but with the expanded quadrate posterior end of septum completely
filling the posterior cavity, or nearly so, as shown in pi. 6, f. 3.
Length, 6 mm. Length of cephalothorax, 3 mm. Length of tib. + pat. I, 2.S mm.;
of tib. -f- pat. IV, 3 mm.
Type— M. C. Z. 356. Claremont.
lb Journal of Entomology and Zoology
Pardosa lifsl<erflta sp. nov.
Ffmalf — Carapace wilh broad side stripes and narrower siipramarginal stripes of
chocolate-brown color, ibe marginal lines black, the eye region also dark; iniddorsal
yellow stripe widest just caudad of eyes with anterior margin straight, from there
narrowing caudad, divided anteriorly by a fine median longitudinal black line, the
stripe only obscurely indicated between eyes. Sternum black. Legs with femora
longitudinally streaked above with black, the joints not annulate or only in part very
vaguely so. Abdomen with a yellow basal mark above, this narrow and widening
caudad; this mark is edged wilh black and is followed behind by several black
chevron lines; venter light, wilh no dark markings. Anterior row of eyes straight or
nearly so, median eyes their diameter or a little mor eapart, about half as far from
the laterals. Eyes of second row once and a half or more their diameter apart. The
epigynum seems clearly distinctive in form. See pi. 6, f. +.
Length, 7 mm. Length of cephalothorax, 3.5 mm. Length of lib. + pat. I, 3.2
mm.; of tib. -f pat. IV, 3.5 mm.
Type— M. C. Z. 392.
Montana: W. M. Mann, collector.
Pomona College, Claremont, California
PLATE 1
Fig. 1. Left male palpus, ectal view, of SemcsniJrs hesfera, sp. nov.
Fig. 2. Left male palpus, ectal view, of Parauximus larJatus sp. nov.
Fig. 3. Epigynum, not fully adult, of Auximiis pallrsii-iu sp. nov.
Fig. 4. Epigynum of Auximus latescens sp. nov.
Fig. 5. Palpus of Aiiximus latescens sp. nov.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Claremont.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. L
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. L
Fig. 2.
Fig. 3
Fig. 4
Fig. 5
Fig. 6
PLATE 2
Left male palpus, ectal view, of Plcitrcurys suprenans sp. nov.
Epigynum of Drassodes celes sp. nov.
Epigynum of Scoinphaeus volunlarius sp. nov.
Epigynum of Herpyllus plus sp. nov.
Epigynum of Zelotes taiho sp. nov.
Left male palpus, ectal view, of Zelnles irritans sp. nov.
PLATE 3
Epigynum of Zeloles //ynethus sp. nov.
Chelicera, ectal view, of Psilochorus caUjorriue sp. nov.
Left male palpus, ectal view, of the same.
Epigynum of Lithyphantes mimoides sp nov.
Epigynum of Teutana grossa (C. Koch), a species not uncommon
Epigynum of Agelena paeifica, var. Claremont.
Eqigynum of Agelena californica. var. Claremont.
Epigynum of Dutyna mians sp. nov.
PLATE 4
Left male palpus, ectal view, of Agelena ma sp. nov.
Epigynum of Olios se/iistus sp. nov.
Left male palpus, ectal view, of Olios schistus sp. nov.
Male palpus, left ectal view, of Anypluena crehrispina sp. nov.
Epigynum of Anyplnena zina sp. nov.
PLATE 5
Right male palpus, ectal view, of Anyplucna mens sp. nov.
Epigynum of Anyp/iiena innirsa sp. nov.
Epigynum of Anyp/nena mundella sp. nov.
Left male palpus, ectal view, of the same.
Epigynum of Anaclieinmis sober sp. nov.
Epigynum of Anachemmis dnlichopus sp. nov.
Right male palpus of the same.
PLATE 6
Epigynum of \amopsiliis pleliis sp. nov.
Epigynum of Lyeosa ferrieulosa sp. nov.
Epigynum of Pardosa tuoba sp. nov.
Epigynum of Pardosa hesperella sp. nov.
Epigynum of Tlianatus retentus sp. nov.
Epigynum of Arariea gosogana sp. nov.
ri.ATK I
J
PLATE VI
Centipedes and Millepedes from Near
Claremont
Most of the specimens were collected during the past few years. All hut the
Scutigera were determined by Or. R. \'. Chamberlin.
CENTIPEDES
SCOLOPENDROMORPH.V
SiolafienJra folymurplin Wood. The largest local form and one of the most
common.
Ulocryptups ijrtnilis (Wood). A smaller form which is also common.
GEOPHII.OMnRPIIA
CtinopoJes limalui (Wood). A species more commonly listed under Mecistoce-
phalus sens. lat.
Sydunguis healhi ialalin,r (Chamb. )
Linot.rnia Ltvipes (Wood). This bright red species is found especially in the
mountains.
Gfofi/iiluj rubfns Say.
Geophilu! rfynans Chamb. Very common.
Arenophilus bipunilicffs (Wood).
Talyuna uniJenlalis (Meinerl).
T. ilarfmontui Chamb.
Tabipliilus rfx Chamb.
Solobius l,rni<ipsis (Wood). A long species witli 129 to 149 pairs of legs.
Gosiphiluj bakfri Chamb.
Gosphilus laticeps (Wood).
SC.TICERO.MORPHA
Siulit/era forceps Raf. From houses.
I.ITIIOBIOMORPHA
l.amyttet pinampus Chamb.
Etho polys xiinti (Wood).
Gosibius pauciJrns (Wood). I'ommon.
Arebius elysianus Chamb.
Solhembius nampus Chamb.
Pokabius clavigerens Chamb.
MILLEPEDES
The species described by Chamberlin were described in the Proceedings of the
Biological Society of Washnigton in December. I91S, Vol. 31, pp. 165-170.
Parajulus jurcifrr Hag.
Tylobolus elarrmonlui Chamb.
Hillonius puluihrus Chamb.
ttuprthnlus (atilorniius Chamb.
./. parvus Chamb.
Spiders from the Claremont-Laguna
Region
The following is a list of spiders collected during the past few vears. All the
determinations were made for us by Ur. R. V. Chamberlin. None of the new species
recently described by Chamberlin are included in this list.
AVICULARIID^
Bothriocyrtum calijornicum (Camb.)
Ulborid.'E
Uloborus uiliforiiiiiis Bks. Uplands, 1200 ft. Nicholson.
DiCTVNID.t
D'ulyna ,ai,aiula Bks.
SCVTODID.^
Plettreurys eastanea Sim.
DR.^SSID^
Zelolcs maiulata Bks.
llerpyllus 'vatidus Bks.
//. antjnstus Bks.
Pholcid.'e
I'hysoiytlus globosus Tac. Uplands Nicolson.
Pholius plialangioides Fuessl.
ACEI.ENID.E
.^getena pacifica Bks.
A. catifornica Bks. Claremont and interior of C'atalina.
A. na-via Hentz. Claremont and Avalon, C'atalina.
Tegcnaria domestica Clerck.
T. californica Bks.
Chorizomma itiliforrm a Simon.
Ll.VVPHIID.'E
Linphia Sp.
Arciopid.e
Cyclosa loniia Pallas. Cucamonga Mt. 4500 to 5500 ft. Johnston.
Aranea miniata VValck.
A. (urcurbitina Clerck.
A. marmorea Clerck.
A. angulata Chick.
Metargiope trijasciata Forsk.
Zilla .\-nolala Clerck.
Tetragnntha labonnsa Hentz.
THERIDIID.JE
Teutana grossa C. Koch.
Latrodectus mnrians Fabr.
T/ieridion lepidciriorum Koch.
Tho.viisid.'e
Thanatus coloradetisis Keys.
Xysiicus rnlifornicus Keys.
X. ferox Hentz.
Misumena vatia Clerck.
Misiimenuides aleatorius Hentz.
Al'tsiimenops asperalus Hentz.
Philodrnmus pcriiix Black.
P. moestus Bks.
Cl.UBIONID.E
Cliiracatithuim uwhisutn Hentz.
TrafJielas tranquilla Hentz. Claremont and Catalina.
Caslianeira pacifica Bks.
Gayenna Juv.
26 Journal of Entomology and Zoology
Anyphocna Juv.
I.vcosio.t
Lycosa kodiii Keys. Clarcmonl and Ontario Mt. 6000 to 7000 feet.
Lycosa, near carolinrnsis, not quite mature.
ParJosa Jlernalis Thorell.
P. lafiiJidna Em.
/'. catijornica Keys.
ParJosa sp. Calallna interior. Claremoni.
Central Nervous System of Mytilus
Californianus
\MI.I,1AM A. mi.TOS'
The cerebral ganglia are well separated from each other, but the smallest of the
three groups of ganglia in the nervous system. A large cephalic branch goes to the
palps, smaller lateral ones supply adjacent parts in the mouth region. No attempt
was made at this time to follow peripheral branches very far.
The single mass of the pedal ganglion may be easily seen to be composed of a
right and a left half. The pedal connections with the main trunk are somewhat
smaller than the long connectives and hardly larger than some of the other branches
of the ganglion, notably the large lateral caudal branches. The caudal branches
of the pedal are chiefiy three on each side, the more lateral being very large and the
medial the smallest. They penetrate and supply the foot and viscera.
The visceral ganglia are more widely separated than the cerebral and much
larger. The large caudal branches pass over the posetrior adductor muscles to be-
come supplied to the muscle and to the mantle. The smaller lateral branch runs
nut laterally dividing soon into two to supply the gills.
In tlie cerebral ganglion the fibers form a broad connection across the middle
line. There are a few cells along the course of the commissure. Nerve cells are
found inclosing the central fibrous mass. The cells are three or four layers thick except
at certain places where there are none. The cells are of several sorts: First — those
that stain deeply with hematoxylin. Some of these may be neuroglia cells, they are
rather small cells and some seem to have very little cell body; second — large cells
with clear protoplasm with distinct fibrillar structure. The processes of these are
long, in some cases may be followed for some distance; third — there are some very
small cells that do not stain deeply. These may some of them be neuroglia cells,
others may be nerve cells in some special physiological condition.
In the fibrous areas of the ganglia, larger and smaller strands are evident, with
only a few cells in central portions.
The cell areas about the cerebral ganglion differs slightlv at various points,
but there is no marked massing into groups.
The pedal ganglion, like the cerebral, has a central fibrous core covered with a
rather even mass of cells of large and small size, but the cephalic and caudal regions
have the thicker masses of cells. The ventral sides of the pedal has more cells than
tlie dorsal. Two sides of the ganglion are well marked from each other, although
broadly connected by fibers.
The large visceral ganglia are more complicated in structure than the others,
but a similar general arrangement of cells is found.
The peripheral distribution of nerves was not followed at this time. For the
general anatomy of bivalves one of the most recent works gives a detailed account
of peripheral distribution in a bivalve:
Splittstosser, P., 1913.
Zur Morphologic des Nervensystems von Anodonta cellensis Schrot. Zeit. f. wiss.
zool. Bd. CIV 3 heft.
(Conliihuliini from the Zoolofi'n/il lahoratnry nj I'umiiita Ciillfije.)
28 Jdiirn.il (/t 1 iitdinology :ind Zoolog\-
Explanation of Figures
Fig. 1. Cemral ganglia of Myiiliis. The cerehral ganglia are above in the fig-
ure. X6.
Fig. 2. Longitudinal section of the cerebral ganglion of Mylilus. The connec-
tive end is down. X70.
Fig. 3. Cross section of the cerebral ganglion of Mytilus. The dorsal side
is up. X70.
Fig. 4. Cross section of the pedal ganglion of \fvtilus. The dorsal side is up.
X70.
~N~^
#
X,-
V )1
Notes on the Sipiinculida of l^aguna
Beach
RALPH V. CHAMIIERI.IX
The followiiii; notes ami provisional diagnoses are based upon a collection
of Sipunculids made hv Prof. Hilton at Laguna Beach in 1917. Six species are
represented.
SIPUNCULUS NUDUS LIXN.
One specimen of this widespread species was taken on the sand Hats at low tide
in August, Balboa. As preserved it is 26 cms. long. Attachments of nephridia and
retractor muscles normal. The usual 31-32 longitudinal muscle bands. The body
appears to be pigmented to some extent, as has also been recorded for a specimen from
the Malacca region described as dark brown (Selenka), and one from Key West
(Cierould). The species is known from various parts of the Atlantic along both shores,
from the Mediterranean, .Adriatic and Red seas, Malacca, Bismarck Archipelago,
Philippines and Japan.
r-HVSCOSOMA ACA.SSIZII ( KEFKRSTEIN)
In the collection are six specimens of this well-marked species taken at low tide
on sand flats at Balboa and I.aguna Beach. The species is otherwise known to occur
along the California coast (e.g., at Monterey Bay, Mendocino, San Francisco, Crescent
City) and northward (Vancouver Id.) as well as southward (Puntarena, Panama).
It has also been taken at Loyalty Is., Ceylon, Laccadives and Maldives, and in the
Atlantic on the coasts of the I'nited States and .Africa.
DENDROSTOMA ZOSTERICOI.A SP. XOV.
This species belongs to the group with but two retractor muscles. These have
thifir origins in the posterior third of the body (in the type about 10 mm. from caudal
end) and are well developed ihroi'ghout and free to near insertions. The nephridia
open at the level of the anrs and are elongate and free. Contractile tube with numer-
ous caeca. 'I'entacles strongly dendritic or arborescent, the terminal branches numerous.
Introvert wholly lacking hooks. Skin brown or in part grey, set off into numerous
arras, these transversely elongate in middle region, in the posterior more rectangular
and longitudinally elongate. Kntire body and introvert, excepting a short area proximad
of tentacles, bearing moderately numerous, small, dark and rounded elevations which do
not rise into true papillae; usually one of these to each cuiicular area; of uniform size
and abiniiance throughout. Body typically narrowed at both ends, fusiform. 35 mm.
long behind anus anil about 15 mm. to distal end of introvert.
Taken in eel-grass in September, l'>17.
Type— M. C. Z. 2, I XI.
OENDRIISroM \ MVTHElTA SP. NOV.
The type of this species was found in eel-grass (Zostera) in the same original
lot with the types of the preceding species. It is a smaller species of obviously different
form. The body is widest at the posterior end, followed by a much longer, narrower,
Pomona College, Claremont. California 31
subcylindrical portion extending to the still narrower introvert proper. The species is
like zostericota in laclving hoolis on the introvert, but differs obviously in the character
and arrangement of the tubercles. These are similarly small over the general body but
are more closely arranged over the middle region than over the broader posterior one,
while particularly characteristic is a band of abruptly much larger tubercles about the
base of the introvert, distad of which region ihey become again abruptly smaller. The
color is brown. The two retractors are inserted at the anterior end of the broad pos-
terior region and are fused anteriorly, their free portions short. Nephridia free.
Length of body behind anus, 12 mm.; in front of this to base of tentacles, near
8 mm.
Type— M. C. Z. 2, 184.
DENDROSTOMA PYROIDES SP. NOV.
Differing conspicuously from zostericota in form, being broadest at the posterior
end and as a whole subpyriform instead of conspicuously fusiform. It is darker brown
in color. A conspicuous external difference is in having the introvert, or proboscis,
armed on its median portion with numerous, comparatively large, dark hooks, which
are not definitelv seriate. The cuticle in general is covered with numerous small, dark,
rounded elevations which in surface view are circular to slightly elliptic in outline
and are larger in size at the base of the proboscis and at the posterior end of the body
than elsewhere. Tentacles arborescently branched, the terminal branches numerous,
finger-like. The two retractors are stout bands taking their origin in the posterior
third of the body. Contractile tube witli fewer caeca. Nephridia free, opening a little
farther forward than the anus.
Length from anus to caudal end, 17 mm.; from anus to base of tentacles, S mm.
Taken at low tide on Laguna Beach.
Type— M. C. Z. 2, 182.
PHASCOLOSOMA HESPERA SP. NOV.
Somewhat resembling P. procerum in form, but with the proboscis more abruptly
set off from the body and on the average narrower and especially much longer relatively
to the latter. In the type the body proper is 8.5 mm. long, while the proboscis is 52
mm. long, i. e., about six times longer than the body, while in one paratype it is as much
as 7.5 times longer. The body of the type is 2.6 mm. thick and the proboscis half or
less than half this thickness. Body proper pointed at both ends, broadly subfusiform.
The skin at the caudal end of the body is rather thickly studded with papillae, which
are disally flat and dark colored over a pale and often constricted base. The papillae
rapidly become fewer and more widely scattered over the middle and anterior regions
of the bodv and over the proboscis, and at the same time become decidedly smaller and
are often borne singly on low, rounded elevations; on the proboscis th epapillae are
tvpically colorless. The two retractor muscles in the type have their origins in the
anterior part of the body.
The type was secured in sand at Balboa, December 26, 1917. Paratyoes from eel-
grass on Laguna Beach, September, 1917.
Type— M. C. Z. 2, 185.
U NOV 1? 1939 i^l
Sine
VOLUME TWELVE NUMBER TWO
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
JUNE, 1920
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT o/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
A Study of the Food Habits of the Ithacan Species of Anura
During Transformation — Philip A. Munz 33
The Central Nervous System of Three Bivalves — WilUam A.
Hilton 57
Journal of Entomology and Zoology
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Tin: ,I<iri:NAi. (ik Rnthmology and Zoology
William A. Hilton, Editor
Clai'pmnnt, (^ilitorniii. 1'. S. A
A Study of the Food Habits of the Ithacan
Species of Anura During
Transformation*
Philip A. Munz of Pomona College, Claremont, California
In recent years almost as much interest has been attached to the study of the
habits of animals and to the relation to the environment as to the structure and
classification. Naturally enough the food-habits are among those that can be most
profitably studied; as an example I have to cite only the work of Professor S. A.
Forbes of the University of Illinois on the food of fresh-water fishes. His results,
embodied in a series of papers published by the Illinois State Laboratory of Natural
History, have been very suggestive and an inspiration to investigation in allied
groups. When, therefore, a study of the food-habits of various species of the Anura
during their transformation period was suggested to me, it was gladly taken as a
subject of some promise.
The purpose of the investigation was to learn something more definite than was
alreadv known concerning the food before and after transformation and to see how
the change from the one kind to the other came about. It was thought that such
knowledge might aid to some extent in frog-culture during this rather critical period
of a frog's development.
The problem was undertaken with the advice and criticism of Doctor A. H.
Wright of Cornell University. To him my sincere gratitude is hereby given, not
only for his suggestion of the problem and for his help in carrying it out, but for
the abundant material which he so generously put at my disposal and which was
the result of much careful collecting on his part.
Largely because of the work he had been doing at Ithaca during the last ten
years, material was available for all eight species of Anura occurring in the Cayuga
Lake Basin of New York state. In all, 586 specimens were dissected, giving a
fairly representative series for each of the species which were as follows:
Rana catcsbeiana Shaw. The Bullfrog.
Rana clamitans Latreille. The Green-frog.
Rana sylvatica Le Conte. The Wood-frog.
Rana palustris Le Conte. The Pickerel-frog.
Rana pipiens Schreber. The Leopard- or Meadow-frog.
llyla crucifer Wied. The Peeper.
Ilyla versicolor Le Conte. The Tree-toad.
Biifo americaniis Holbrook. The Common Toad.
METHODS
As specimens were collected in the field they were immediately killed, usually in
formalin, in order that digestion would immediately cease. Each lot that was col-
A contribution from the Zoological Laboratory of Cornell University of Ithaca, New York.
34 J(jurn;il of Entomology and Zoology
lectcd was kept separate and the data as to its time and place of collection were
recorded. This data can be found under the discussion of the individual species.
My part in the collecting was during the season of 1916, and during the following
fall and winter the material was examined.
As complete information as possible was secured as to the stage of transforma-
tion; the length of the tail, condition of the mouth, length of the alimentary canal,
its differentiation into stomach, small and large intestines, and the development of
the front legs — all these facts were noted. Unfortunately at first the desirability of
so complete a record was not realized and the bullfrog, which was the first species
studied, did not receive as full treatment as those taken up later.
After the alimentary canal had been removed and its length had been measured,
the contents were removed and identified. In many cases, prhaps a more exact
determination of the forms found could have been carried out by specialists, but the
kind of food rather than the exact species seemed the essential thing. For this reason
a not very serious attempt was made below the identification to family, especially
where digestion had proceeded to any extent.
WORK PREVIOUSLY DONE
Some very thorough investigation has been carried out on the food of the adults
of several species, in America the most notable being that of Kirkland on the Ameri-
can toad, and of Drake on the meadow-frog. Kirkland' in an examination of 149
load stomachs which had been collected in a number of situations, found that by
bulk 98'f of the food examined was animal, that 77'/r was made up of insects and,
that of the insect food 11' I was of beneficial forms, 22'/ neutral and bl''i injurious.
He made note, too, of the fact that the toad feeds largely at night and that in a single
twenty-four hours it can fill its stomach to its complete capacity four limes. He said,
too, that the toad takes only living and moving forms; this fact is one repeated
by other observers for other species of the Anura and agrees with my own results.
As an example of the evident attractive power a moving object has for an .\nuran I
may mention a specimen in the Museum of the University of Denver. It is a toad
probably of the Woodhouse variety, which was brought in dead and dried up and
with the lip of a turkey wing projecting from the mouth. It had evidently been
attracted by the small bunch of feathers being blown about and, having swallowed a
part was not able to finish the process nor to disgorge because of the barbs of ihe
feathers catching in Ihe throat.
From the tables given by Kirkland one is led to infer that practically no a(|ualic
forms enter into the load's diet, a not very surprising fact when its terrestrial habits
are remembered. Since many a(|ualic insects are attracted lo electric lights, it is
evident that a load or frog feeding under the corner arc light can secure such forms
without ever approaching water.
'Kirkl.in.1, .\. II.. 1897. Habits, food .nn.l icmoniic v.ilvic of the .\nicrican toa.l. (Bull. A6
of Hatch hxp. Station of the Mass. ARric. College. 1904.) Usefulness of llic American toad.
(Karmen Bull. 196, U. S. llcpt. of Agricullurr.)
Pomona College, Clareniont, California 35
In addition to the work of Kirkland some other work has been done by Garman"
on the toad in Illinois and Kentucky, and has had in large part the same results,
showing that many harmful insects are destroyed.
The work of Drake' mentioned above and dealing with the meadow-frog covered
an examination of 209 stomachs. Mr. Drake says, "All the evidence indicates that
the presence of substances other than those of an animal nature is merely incidental,
and due to the mode and conditions of feeding," and "Nothing can be more natural,
since the frog captures the greater part of its prey on the ground by means of its
tongue, than that a small amount of foreign substances should be swept into the
mouth along with the animals upon which it feeds. The frog's food consists of
mollusks, crustaceans, myriapods, spiders and insects; in fact any sort of living
creature is acceptable to it as both sense of taste and of smell are apparently obtuse."
In his work at Saranac Inn in the Adirondacks, which is reported in the "May-
flies and Midges of New York," Professor Needham' spent some time in a study of
the summer food of the bullfrog and in his report gives the contents of 16 stomachs.
Of the 164 animals found 139 were insects, 18 were snails, three crustaceans, three
spiders and two vertebrates, one bullfrog tadpole, and one meadow mouse. This
assemblage differs largely from the food of the toad in that many aquatic forms
are reported, some of which must have been taken under water, for example the
nymph of the mayfly Siphliirus alternatus Say, which Professor Needham says never
comes to the surface except for transformation. Other forms were the Rittiii tadpole,
Anax nymph, chironomid larvae, and a small copepod and some aquatic snails; while
the water-striders, soldier-fiy larvae (Stratiorayidae), gnat pupa: and transforming
caddisflies taken were probably secured at the surface of the water.
The few others who have studied the food of the bullfrog have likewise found
many vertebrates eaten. Brakeley" in twelve bullfrogs dissected found one mouse,
one young bird, one frog, two toads, two carp, six mud-fish, one mud-turtle, besides of
course, many insects and other invertebrates. Dyche's report" in 1914 on the con-
tents of 30 bullfrog stomachs substantiates previous reports on the greed and rapa-
ciousness of this species. He found that one specimen thirteen inches long had
swallowed another about ten inches long and cites many other cases of cannibalism.
Fourteen of his 30 species contained 32 fish, otherwise the diet was made up largely
of crayfish, other crustaceans, insects, spiders and snails.
Brief mention of the feeding habits of the species of Anura is made by Miss
Dickerson' who has evidently based some of her account on personal observation.
Of especial interest here are remarks on the food of tadpoles. Concerning the tad-
pole of the toad she says, "These mouths are provided with horny jaws for scraping
the tiny plants from their supports and for biting off the delicate ends of larger
plants." In another connection, "The tadpoles of the wood-frog eat not only the
green jelly mass from which they themselves hatch, but also the soft green spheres
within the jelly masses vacated by young salamanders. Like other tadpoles, they
=Garman, II., 1901. The food of the toad. (Bull. 91, Agric. Exp. Station of Kentucky.)
'Drake, C. .1., 1904. The food of Rana pipiens Shreber. (Ohio Naturalist. 14:257-269.)
*Needham, J. G., 1905. The summer food of the bullfrog (Rana catesbeiana Shaw) at
Saranac Inn. (Bull. 86, New York State Museum.)
=Brakeley, J. H., 1885. Notes on carp and frog-culture. (Bull. U. S. Fish Com., 5:209-213.)
'Dyche, L. L., 1914. Ponds, pond fish, and pond fish cuhure. (Bull. No. 1 of Uept. of
Fish and Game, Kansas.)
'Dickerson, Mary C. The Frog Book. (Doubleday Page and Co.)
36 Journal of Entomology and Zoology
act as scavengers by greedily devouring all dead animal matter of tlie pond." For
the bullfrog tadpole she says, "The mouth is well fitted to bite the delicate ends of
leaves and stems, or to scrape off the tender green or brown plants from sticks and
stones. It is equally adapted for eating animal food. The bullfrog tadpole (like
that of the green-frog and that of the wood-frog), is especially fond of any animal
food available. Thus these tadpoles act as scavengers and dispose of dead fish or
dead tadpoles even, that otherwise would become a menace to the living creatures
of the pond."
Her remarks on the food of the adults bear out the statements made by other
workers and show further that the bullfrog and the green-frog are the most aquatic
of the eastern frogs, that the meadow- and pickerel-frogs are less so, while the
wood-frog, peeper, tree-toad and common toad are even less inclined to be found
in the water. The food is therefore expected to vary accordingly.
Her observations on moulting the skin are also of interest. "The green-frog
moults the skin four or more times each year. H the frog is out of the water when
the moulting takes place, the process is like that of the American toad and of the
leopard-frog, and the skin is swallowed. If the moulting takes place in the water
the skin may float off in large patches and is not eaten."
From this short review of the literature in regard to the food of the Anura
of this country, it can readily be seen that there are many statments as to the food
of the adults and some remarks are even made as to that of the tadpole, but I
have found nothing as to food during transformation, except that the tail is absorbed
by phagocytic action and is used. It shall therefore be my attempt in the following
pages to take up in more detail the food of the adult tadpoles, of the transforming
ones, and of the young frogs.
GENERAL DISCI SSION OF TRANSFORMATION
One of the first evidences externally of transformation and the one most greatly
affecting the food situation is the shedding of the horny plates characteristic of the
tadpole's mouth. .After this takes place the alimentary canal decreases to one-tenth
or one-twelfth of its larval length, and at the same time becomes differentiated into
distinct portions. The larval digestive tube is merely a long tube, that of the young
frog has a widened and thick muscular-walled stomach, a long narrow small intes-
tine, and a much shorter but likewise thin-walled large intestine. In the specimens
dissected the longest larval alimentary tract found in the bullfrog was 1070 mm.,
the average after decrease in length was from 85 to 95 mm.; for the green-frog the
two measurements were 450 and 50 to 60 respectively, for the wood-frog JOO and
18 to 2(1, for the pickerel-frog 400 and 30 to 35, for the meadow-frog 530 and 30
to 40, for the peeper 88 and 10 to 15, for the tree-frog 170 and 15 to 18, and for the
toad 110 and 10 to 15. While all this change is going on the mouth is gradually
increasing in size to one many times larger than the tadpole mouth. Obviously while
the mouth is still very small it is scarcely capable of taking in any food, its horny
plates having been lost. In order that the materials eaten may be swallowed whole
and not nibbled off, the mouth has to be considerably enlarged. It is not strange
then, to find that without exception all eight species go through a period of fasting
as far as taking food from the outside is concerned. An examination of the data
Pomona College, Claremont, California 37
given under the separate species will show that many of the individuals studied, and
practically all of these in which the mouth was enlarging, contained no food. It
seems hardly necessary to say in this connection that the absorption must make
possible this period of fasting.
It must take some little time for the food to pass through the long alimentary
canal of the lar\a, since it was quite noticeable that in many of those examined in
which the mouth plates liad recently been shed, the caudal portion still contained the
mud and other contents typical of the larva and which had probably been taken into
the body when the mouth plates were still in place.
The length of the body without considering the tail, remains almost unchanged
during transformation, but the rotund aspect of the tadpole gives way to the flatter
and more angular one of the young frog. The appearance of the front legs, the left
one coming out through the spiracle and the right one breaking through the skin,
as well as the shrinking of the tail are further indications of the progress made in
transformation. Usually the tail is almost gone before feeding as a carnivor begins.
It is to be noticed that in the data given for the bullfrog almost every trans-
forming individual is reported as having swallowed some of its own cast epidermis.
This is true to a lesser degree of the other species, probably because the mucli smaller
size of such species as the tree-toad and the toad makes the rcognition of epidermis
in the alimentary tract of preserved specimens more uncertain. The frequent occur-
rence, one might say almost universal occurrence, of epidermis in transforming
individuals must mean frequent moulting. Doubtless this is true, especially of the
tail, which shrinks rapidly and might naturally shed its epidermis. That the shedding
of the skin takes place in the water is evidenced by the threads of Spirot/yra and
Zygnema often wrapped up in it as if during seizing and swallowing.
The discussion of transformation may be concluded, then, by saying tliat it is
accompanied by a period of fasting during which time, in all eight species, the food-
getting and food-assimilating apparatus is rebuilt and changed from one suitable
to a form largel\ herbivorous and at least feeding only on dead animal material,
to one wliicli will permit of the predaceous and carnivorous habits of a frog or toad.
PRESENTATION OF DATA FOR THE SPECIMENS EXAMINED
In the following pages lists are given of the specimens dissected with data show-
ing the degree of transformation and the contents of the alimentary canal. The
word "stomach-content" is not sufficent here; for in many cases the stomach was
almost empty while the large intestine contained large amounts of food; in the case
of insects, passage through the digestive tract in this way had not sufficiently changed
many specimens to make it impossible to identify them to family at least. In the
data given under each species, "No." refers to the number of the specimen, "Body"
to the length in millimeters, measuring from the tip of the head to the base of the
tail; "Tail" to the length of the tail in millimeters, "Mouth" to the condition of the
mouth, whether that of the tadpole with the horny plates or with these shed, or with
the mouth enlarged; "Ali. Can." to the length in millimeters and to the condition
of the alimentary canal; "Fore Legs" to the presence or absence of the front legs;
"Lot" to the lot to which the particular specimen belonged, and "Food" to the
material found in the digestive tract.
38
Journal of Entomology and Zoolog)'
Rana calesbeiana Shaw. The Bullfrog.
Total of 104 specimens. Lot 1, Dr. A. H. Wright and Dr. A. L. Leathers,
Wood's Hole, Mass., July 16, 1909. Lot 2, Wright and myself, July 10, 1916, Dwyer's
Pond, Ithaca; lot 3 same, but on July 17. Lot 4, Dr. Wright, July 7, 1911, Beaver
Brook Mill Pond, Ithaca.
Table 1. Data for Rana catesbeiana
No. Body Tail
I -18 87
.Mouth .\li. Can.
tadpole 1070. ladpult-
45 95 tadpole 980, tadpole
changing 340, stem, small
tadpole 140. stum, small
changing 100. stom. & int.
small 115 ••
1-"U01)
iMud with Mcridion, Cymbclla, Navlc-
ula, Diatoma, Frustulia, Pinnularia,
Oscillatoria
Mud with Epitheniia, Navicula. Pinnu-
laria. Diatoma, Synedra. Nitzchia,
Cymbella, Mcridion, Ulothrix, Spiro-
g>-ra, Oscillatoria
Green algal threads
Greenish material, not identiBable
Epidermis, Spirogyra, Zygnema
Epidermis, Spirogyra
Nothing
80
I Epidermis.
Spirogyra
80
165 •• '• II
Epidermis
70
.Nothing
70
Nothing
70
Nothing
68
110. Slum. &■ int.
Epidermis
68
Epidermis.
piece of plant tissi
65
100 ••
Epidermis
65
Epidermis.
Insecla 1
64
Epidermis
60 • "
100. stom. & int.
Epidermis
60
Epidermis?
60
stom. & int.
Epidermis?
58
94
Epidermis
57
85
Epidermis
55
95 '
Epidermis
55
97 •• " ;•
Epidermis.
some plant tissue
52
Nothing
52
78, stom. & int. "
Epidermis
52
88
Epidermis
So
85 ■• ■• '•
Epidermis
50
Nothing
50
Epidermis
50
slom. & int.
Epidermis'
48
Epidermis
48
Epidermis
47
slom. & int.
Epidermis
46
Epidermis
46
stom. & int. •■
Epidermis'
45
Epidermis
45
115. stom. & int.
Epidermis
45
Epidermis
45
stom. & int.
Epidermis
Ai
95
Epidermis.
sand, mosslcaves
43
96 •• •■_ II
Epidermis
43
Epidermis
40
90
Epidermis
40
82 •• " II
I'.pidermis.
some plant tissue
40
El>idermis
40
^i. stom. & int.
Epiilcrmis.
Collembola 1
40
Nothing
.18
stom. & int. "
Epidermis
35
95
Nothing
35
85
Epidermis
35
Eltidermis
35
stom. & int.
Nothing
32
Epidermis
32
stom. & int.
Epidermis?
Copcpoda 2
30
l-'iiidermis
28
95, stom. & int.
Epidermis.
plant tissue
28
Epidermis,
Copepoda 1. Colic
27
«« ft M
Nothing
Pomona College. Claremont, California
39
Table 1. Data for Rana catesbeiana — Continued
No. Body Tail Mouth
93. stom. & in.t
90, stom. & int.
Epidermis
Epidermis
Epidermis,
Diptera
Xothing
Epidermis
Epidermis
I'hiloscia (
Epidermis
ella
Epid
Epid
Epid
Epid
Xothing
Epiderm
Epiderm
Epiderm
Epidermis?
Lestes vigila-t (2
Staphvlinidae 1,
Difflugia 3. Hyd:
CoUcmhoIa 1. Capsiilae 1,
green
Ciirculionidae I
Mv
Epidermis
Collembol;
Epi.lcrmis
-\nhididi
Epidenr
Collembol
dae 1,
Spirogyra
Coleopte
2, Staphylini-
Laccophilus 1. Dascyllidae 1
larva. Formicidac 1, egg I. achenc
of Scirpus
Epidermis, unidentifiable material
Collembola I. plant fiber
Epidermis, unidentifiable material
Nothing
.\carina several. Lestes nymph
Xothing identifiable
Collembola 1. Cercopidae 1
Acarina 1. Collembola 1. Cercopidae 1.
-Agromyzidae 1. .-\nthonomus (Cnr-
culionidae) 1. .-\phodius 1. bits of
Sphagnum
Coenagrionidae 1. Carabidae 1
Gerridae 1. F.epidoptera 1 larva. Clado-
phora. Spirogyra
Coenagrioninae 1. nymph
Copepoda 1. Agromyzidae 1. Elateridae
Elateridae 1. young Rana catesbeiana,
grass seed
Clubionidae (Araneida) 2. Oribatidae 1.
Collembola 3. Creniphilus I. Coleop-
tera 1. Sphagnum, twig
Oribatidae many. Collembola many. Pa-
norpidae. 1. Cercopidae 1. .\gromy-
zidae 1. Coleoptera 1. Hvdrophilidae
1 larva
Aphididae several. Agromyzidae 2
Nauplius many. Copepoda several. Col-
lembola 1. Coenagrioninae 1. Coleop*
tera 1. Elateridae 1. Curculionidae 1.
Hymenoptera 1. Aphididae 1. Bidens
seed. Sphagnum lea
ves, plant fibers
ifflugia. Copepoda m
any. .^carina sev-
eral. Coenagrionin
ae 1. Zygoptera
nymph. Psocidae
1. Hemiptera I.
Coleoptera 1. Chrvs
omeelidae 1. Car-
abidae. Curculionida
s 1. Hydrophilidae
1. young Bufo. mo
s. thistle-down
entatomidae 1. Rhyn
cophora 2, Hydro-
philidae 1. Apida
1, much dirt.
straw, moss-leaves.
seeds, pieces of
chitin
40
Journal of Entomolog>' and Zoology
Rana clamilans Latreille. The Green-frog.
Total of 87 specimens. Lot 1, Crystal Beach, Canada, June, 1914; lot 2, Casca-
dilla Creek, Ithaca, July 22, 1907; lot 3, Bool's Backwater Ithaca, June 30, 1906; lot
4, Slaughter House Ponds, Ithaca, June 20, 1906; lot 5, same, June 10, 1907; lot 6,
same, June 27, 1911; lot 7, same, June 30, 1911; lot 8, Dvvyer's Pond, Ithaca, date
not given; lot 9, Slaughter House Ponds, June 29, 1907; lot 10, same, June 29, 1907;
lot 11, Cascadilla Ponds, Ithaca, July 7, 1916; lot 12, Owyer's Pond, July 10, 1916;
lot 13, Wood's Hole, Mass., July 16, 1909; lot 14, Dwyer's Pond, July 27, 1916; lot 15,
Michigan Hollow Pond, Ithaca, Aug. 5, 1916; lot 16, Biological Station, Ithaca, June
30, 1911; lot 17, Bool's Backwater, Ithaca, Sept., 1912; lot 18, Ithaca, June 21, 1915.
Lots 1-10, 16, 17, 18 collected by Dr. Wright; lot 13 by Dr. Wright and Dr. A. L.
Leathers; lots 11 12, 14, IS by Dr. Wright and myself.
Table 2. Data for Rana clamitans
.S'o.
Body
Tail
Mouth
.\li
. Can
I-
ord.-gs
l.o
'
34
63
tadpole
tadpo
le.
230
one
7
-
32
60
tadpok
290
none
7
3
29
60
tadpole
stomach,
110
present
11
•I
29
Si
160
none
12
^
30
.S3
..
ta.l|)o
Ic.
440
none
12
6
28
53
300
none
12
~
32
52
..
450
none
4
8
28
$0
stomach.
250
9
9
29
60
110
present
11
10
32
45
l.i '
ilf.
440
none
7
31
55
chaneinic
stomach.
120
present
16
30
53
small
55
U
31
53
••
94
16
33
52
tadpole
tailpii
>ic.
90
12
27
5!
small
90
12
31
SO
slom.
&int
. 57
II
31
55
small
stomach.
120
16
IR
31
47
**
stnni.
& int
11
19
30
46
55
11
«Ioin. & int. 45
2fl 27 42 small
29 28 40
.15 33 50
J6 J3 32
one-half " *'
onc-foufthslomacb'.
Mom. ft int.
stomnch,
FOOD
Mud with Ivunotia, Diatoma, Navtcula,
Syncdra, Spirogyra, Zygncma, Aoa-
bxna.
Mud with Diatoma, Navicula, Eunotia,
Nitzchia, Synedra, Paramcecium, Zyg-
nema, Cladophora, Spirogyra, Mou-
geotia.
Nothing.
Mud with Oscillatoria. Navicula, Spiro-
I gyra, Nitzchia. Cymbclla, Ccriodaph-
• nia, Kuglena ?
Plant tissue, algal filaments.
Mud with Synedra. Pinnularia, Na-
vicula. Diatoma. (".omphoncina. Nitz-
chia. Oscillatoria. Kuglena ?
Mud
Mud with tadpole teeth. Zygncma, Mi-
crospora, Diatoma, Navicula. Com-
phonema.
Nothing.
Mud with Spirogyra, Mougcotia. Zyg-
ncma. Oscillatoria. Kunotia. Navicula,
Synedra. Tabcllaria. Nitzchia. Melo-
sira. t'oniphoncma. Clostcrium.
Nothing
Nothing
Nothing
Nothing
Nothing
Kpidcrmis
Nothing
Nothing
Kpidcrmis
Kpi<lermis,
Kpidcrmis
Mud with Navicula, Gomphonema, Os*
cillatoria. Spirogyra. cells of pine
wood.
Kpidcrmis, Spirogyra
Nothing
Kpidcrmis
Kpidcrmis ?
Kpitlermis. Spirogyra. Zvgncma
Kpidcrmis ?
Nothing
Kpidcrmis
Kpidcrmis
Kpidcrmis. Spirog>*ra
Hpidemiis ?
Nothing
Nothing
Kpidcrmis
algal fiilamcnts
Pomona College, Claremont, California
41
Table 2. Data for Rana clamitans — Continued
64 42 n
FOOD
, plant tissue
, Spirogyra, Zygnema
I-pide
Upide:
Epide
Epide
Epidermis
Epidermis ?
Epidermis
Epidermis, sand
Epidermis
Agromyzidre 1 , Coleopt
1
. Dytiscidre
Leptidve 2,
Epidermis, Agromyzida
Formicidx 1
Eycosidse 1, Lygaeidae 1, Ceratopogon
larva. Curculionids 2, Braconidae 1
Tipulids 3, Megilla maculata (Coccinel-
lids)
Epidermis, Drassidse 1
Insecta 1
Zygoptera nymph, Leptidx 1, Formi-
cidas 1
Tipulidi' 1
Lycosidw and egg-sac, Scaraba?ids 1,
Formicidffi
Epidermis, Coleoptera 4
Pelecypoda, Agromyzida
1. Fo
licids 1
1, Coleople
Phalangids 1. Tassids 1, dirt and tr,
Coleoptera 1. Staphylinids 1. Diptera I
L,ymnffid.'e 3, Nematoda 3, Araneida 1,
Insecta 1, larva, Capsid^e 1, Jassid:e
1, Lepidoptera larva
Nematoda 1, Panorpids 1, Lepidoptera
larva, Formicids 2, mass of eggs,
fruit of Juncus
Epidermis ?, Dictynus (Araneida),
Tipulidae 1
Trematoda 1, Limnobatidse 1
Libelluline nymph, Hydrophilids 1.
Psilopus (Dolichopodid.-e) 1, Plant
material and sand
Araneia 1, Anisoptera nymph, Curcu-
lionidne 1. Formicidae 1, Myrmicid.-e 1,
Ponerid.T 1
Diplopoda 1, Oniscidae 1. Araneia 1.
1 Jassid.-c 1, Corixidas 1, Heteroneurid?e
1, Diptera !. Eeptidas I. Anthomyidre
1, Curculionidae 3. Ichneumonids
Ostracoda 1, Trematoda 4, Gastronoda 1,
Jassid.-e. Coleoptera 1. Dytiscidae 1.
1 Chrysomelidre 1, Rhyncites bicolor 1,
Ephyhdridse 1. Formicidae
Lymnaea 2, P.orcellio rathkei 6, Ceomet-
rid.-E 1. larva. Carabids 4
Porcellio 1, grass, mud. plant fibers
Coptocychla guttata 1, Diptera larva.
Carabid.-e larva, bits of grass
Porcellio 2. Theridiids 1, Carabid.-e 4.
Mud grass
Epidermis. Capsidas 1, Empididae 1.
Carabidff. Tenebrionidae 1
Araneida 1 , Insecta 1 , Carabidae 1 ,
larva and 1. adult, mud. plant fiber
Hemiptera 1. Diptera 1, Carabid.-e 1
Coleoptera 1, Carabida; 1. Curculionid.-e
Lun
I. Ca
Epiderir
Chrysomelid^
Polvtjra ? (Gastropoda) 1
Oniscids 6, Lvm.Ta 2. Lithobius 1 .
Diptera I. Coleoptera 1
Oniscida? 4, Lumbricidze, Lymn,-ea 1 ,
Argiopid;e 1. Potomogeton leaf
Argiopoidea 1 , Diptera 1 . Empidid-T 2,
Coleoptera 1 . Potamogeton leaf
42 Journal of Entomolog>- and Zoologj'
Table 2. Data for Rana clamitans— Continued
No. Uody Tail Mouih Ali. Can. Fore LfRS Lot FOOD
80 J5 0 '■ ■■ " 86 " 16 Nothing identifiable
81 34 0 •• '■ " 75 " 16 Capsidi 1, Jassida: 1. Uiptera 1, Empi-
dida.* 1, Carabidx 3
82 36 0 r.S '■ 18 I'halangidat 1, Phitenus lineatus 1. Dip-
tcra adult and larga, Carabidx 2,
Kormicidx 1, Salix fruit; epidermis
83 34 0 I 5 " 18 Nothing identifiable
84 36 II >^^ " 18 Phalangid* 1. Carabida: 1, Uiptera 1,
larva and 1 adult. Lcptida: 1
85 37 n 72 " 18 Oniscidx 18 nymphs, Hcmiptera 1, Ccr-
copidx I nymph, Capsidx 1, Diptera
lar\'a, Carabidx 2, Rhyncophora 2.
86 37 0 1 "' '■ 18 Hemiptera 1. Tipulidx 1, Carabidx 1.
Salix fruits 3, straw, unidentifiable
material
8" 37 0 "(' 18 Oniscidx 1, Diplopoda 1, Dolichopodidx
' 1. Coleoptera lar\'a. Formicidx 3 epi-
dermis, Salix fruits 6
Rana sylvalica I,e Conte. The Wood-frog.
Total of 100 specimens. Lot 1, Hamburg, \. Y., July 1, 1907; lot 2, Beehive
Pond, Ithaca, July 32, 1907; lot 3, Cross-road Pond, Iihaca, July 4, 1907; lot +, Bee-
hive Pond, June 2S, 1911; lot 5, Cross-road Pond, Ithaca, July 5, 1907; lot 6, Beehive
Pond, July 31, 1907; lot 7, Beehive Pond, July 8, 1908. Lot 1 collected by Dr. A. A.
Allen of Cornell University; lot 3 by Dr. Wright and Dr. H. D. Reed; the others
by Dr. Wright.
Table 3. Data for Rana sylvatica
No. Body Tail Mouth Ali. Can. Forelegs I,ot , FOOD
1 22 36 tadpole lailpolc, 200 nunc 4 Mud with Cymbella, Navicula, Nitzchia,
Diatoma, Amphora, Fragillaria, Epi-
tliemia, Meridion, Microspora
2 16 28 ■' •■ 65 " 3 Mud with Difflugia, Eudorina, O.scilla-
toria
3 19 2» " stiim.ich, 60 present 4 Mud with Navicula. Pinnularia, Meri-
dion, Stcplianodiscus, Arcclla. Epi-
dermis ? •
4 IS 25 " tadpole, 130 none 2 Mud with Spirogyra, Oscillaloria, Clos-
terium. Navicula
5 18 20
17 13 20
'*
vK.mach.
105
present
Mud with -Vrcclla. Pialoma, Navicula
Synedra, Sccncdesmus
"
sl.mi. &' inl.
28
none
Plant tissue, fibers; .Navicula
*■
slnmach.
IS
Nothing
smnll
37
35
present
Nothing identifiable
Nothing
"
stnm. & int.
28
30
1!
Nothing
Nothing
"
25
Nothing
"
sinmach.
S3
Some mud with Navicula and llialoma
**
slom. & inl.
IS
Nothing
"
" *'
24
Epidermis
"
" "
24
Nothing
"
" •'
18
.Nothing
"
tadpole.
14
••
Mud with Navicula and Fragillaria
"
14
Nothing
"
slom. & int.
211
Hiatonia. Mnugcotia
changing
t.vliHile.
40
Muil with ('■oniphonema and Synedra
<iniall
'<l<>m. & inl.
28
18
Nothing
Nothing iilentifiable
*'
^umiach.
21
none
Plant tissue
tadpole
Mud with Diatoma. Navicula. f.ompho-
nema. Fragillaria
large
!tlomach.
17
present
Nothing
«mall
31
Epidermis
Pomona College, Claremont, California
43
Table 3. Data for Rana sylvatica — Continued
Mouth
one-half
small
of Mougeotia and
FOOD
Nothing
Epidermis
Nothing
Plant tissue and fibers
Epidermis ?, Zygnema,
, bonema, CynibcUa, N
' laria
Epidermis ?
Epidermis ?
Epidermis, Zygnema
Epidermis ?
Epidermis ?
Epidermis ?,
Zygnema
Plant tissue
Epidermis, Zygnema, Mougeotia, Oscil-
latoria
Epidermis ?, Spirogyra, Mougeotia
Nothing
Epidermis ?
Nothing
Nothing
Epidermis, Zygnema
CoUembola 7
Epidermis
Epidermis
Epidermis, Psocids 1
Epidermis ?
Epidermis
Epidermis ?
Epidermis, Zygnema
Epidermis
Epidermis, Psocids 1
Epidermis
Nothing
Planorbis 1, Collembola 1, Chrysomel-
id:e 1, Proctrotrupidre
Asellus 1, Chironomid.-e 1, Proctrotru-
pids 1
Diptera 1
'ith Spirogyra and Eudori
vicula, Oscillatoria
I Chi
1. Corrodenti:
dx 1 lar
Diptera 1.
Hydrophilidre
■'''"'^ 1 larva
3 Coleoptera larva
3 Epidermis
4 Epidermis
2 Acarina 1, Diptera 1
6 Psocids 2, Cynipids 1. Proctrotrupidn?
1 Hydrachnidre 2, Hydrophilidae 1
1 Epbvdridas 1. Chrvsomelida 1
1 Aphidids 1, CurcuHonidx 1, Proctro-
trupidae 1
1 Chironomidae larva, some unidentifiable
material
1 Diptera 1, Hydrophilidae larva
2 Collembola 1. Corredentia 1, plant tissue
2 Collembola 2, Hydrophilids larva. Cyni-
pidae 1
2 Ephydrids 1, Cynipidse 1, unidentifiable
material
5 Araneia 1, Aphididse 1, Diptera 2, plant
tissue
Eygsidse 1, unidentifiable
Insecta la
material
Green alg;
Planorbis
Hete
i 3. Oniscidal. Collembola 3.
ptera 1, Eraconids 1
Collembola 7. Diptera 1, Proctrupidx 1,
Braconidic 1, 4 anthers
Diptera 3. Carex seed
Epidermis, Collembola 1, Hydrophilids
lar\'a, sand, Proctrotrupidae ^ anther
44 Jdiinial of Entomology and Zoology
Tabic 3. Data for Rana sylvatlca — Continued
No. Ilo.ly I ml Monilv Ali. Can. Fore Legs Lot KUUU
85 16 II ■• " 22 " 6 Insccta 1. Jassidx 1, Diptcra I. Chiro-
nomidz 1, Tipulidx 1. jiyiirophilidx 1
86 17 0 -^~ '• Acarina 1. Chironomidx* 4 larva;, l,epi*
iloptera larva
8' IS 0 Diinugia 1, Acarina 1, Collembola 2,
IJiptcra 2 larvx. Phorido: I, seed,
stellate trichomes of plant
88 17 0 J4 6 Jassid.x- nymphs
89 16 II -'3 •• 7 Collembola 3, FulgoridK 1, Slaphylini-
d:u I, Ucpidoptera 2 larva;, Diptera 1
90 17 0 27 " 7 Ciirculioniila; I, Phorida; 1, Braconida;
1. Proctrotrupid.-e 1
91 17 0 •• " 35 " 7 Lycosid:c 1. fhorida; 1, Anthomyid.i; 1,
Staphylinidx, Proctrotrupida; 1, Cyni-
, pida; 1, Chalcididx 1
92 16 0 " •• 22 7 Diplera 1, Chalcidids 1
93 17 0 " '■ 28 " 7 Diptcra 1, Phoridx 1, Chalcididx 1
94 17 0 •• 28 " 7 Araneida 1. Collembola 5, Uiptera 1,
Coleoptera 1, Chalcididx 1
95 15 0 18 " 7 Collembola 4, Diptera 1
96 IS 0 -'5 " 7 Collembola 2. Phoridx 1. Diptera 1,
Ichneumonidx 1. bit of down feather
97 15 (1 :8 •■ 7 l.inyphiidx 1. Kulgoridx 1. Diptera 1.
Colcoplura 1, Hydrophilidx 1, Curcu-
liiMiid.-e 1. insect eggs
98 14 0 J8 •' 7 l.inyphiidx 2. Psocidx 5 nymphs, Dip-
tcra 2, 'Proctrotrupid-x 2
99 15 0 23 " 7 Orilutidx 1. Coleoptera larva, Proctro-
trupidx 1. one empty anther
ino 13 0 23 •■ 7 Psocidx 3 nymphs. Lepidoptera larva.
Diptcra 1. Coleoptera 1. Staphylini-
dx 1
Rtina paluslris Le Cniitc. Tlu' I'ickcrcl-f rog.
Total of 10(1 specimens. Lot 1, Michigan Hollow Pond, Ithaca,- Aug. 5, 1916;
lot 2, Bool's Backwater, Ithaca, Sept. 1, 1912; lot 3, same, July 29, 1907; lot 4, Cross-
roads Pond, Ithaca, Aug. 6, 1907; lot 5, Bool's Backwater, Aug. 6, 1907; lot 6, no
date nor locality. I.ot 1 collected by Dr. Wright and myself; lots 3, 4, 5 liv Hr.
Wright; lots 2 and 6 presumahlv In him.
Table 4. Data for Rana palustris
No. Ilody Tail Mouth Ali. Can, Forelegs I.oi FOOD
1 ■' : tadpole tadpole, 200 none 1 Mud with Mougcotia, Xlcrismopedia.
Microcystis, Scencdcsmus. .Navicula,
I'iniuilaria, lliatonia. Kunotia, Nitz-
chia. Cosmarium, Closteriuni, Pando*
rina
2 25 47 changing ladpnle. 168 " 2 Mud with Navicula, Dialoma, Synedra,
ICpithcmia, Comphonema, Oscillatoria,
.\lougcotia. Spirogyra, Zygncma. Seen-
clcsmns, Clostcrium
3 23 44 tadpole " ISO " 1 Mud with Comphonema, Navicula. Dia-
toma. Pinniilaria. Scenedesmus, Clos-
tetrium, Cosmarium, Merismopcdia
* 24 33 ■' •■ 400 " 5 Mud with Difflugia. Diatoma, Navicula,
5 22 30 changing " 135 one 3 Syneiira, Kunotia. Comphonema,
Nitzchia, Melosira, Scencdcsmus, Kpi*
tlicmia, Spirogyra
Mud with Navicula. Ocdogonium, many
Strongylli<lx, the latter probably para-
sites
6 .^,1 40 sinnll ,i,,ii,.-„h 1 /•(I „„.■ | jfud with Merismopcdia, Oscillatoria.
Closteriuni. Scenedesmus. Cosmarium,
.\avicula. Protozoan parasite
Pomona College, Claieinoiit, California
45
Table 4. Data for Rana palustris — Continued
two-thirds
one-third
one-half
LcRS Lot
FOOD
cscnl 4
Nothing
1
Nothing
1
Mud with Clostcruini, I'aranirecium,
I'leuroc-orcus, plant tissue
1
:• ,>idermis ?
3
l,ittle mud at end of ali. can.
1
'* 1
Nothing
i,',,;,i,.,-i,iic ?
1
i.piuLiniis r
X,. thing
1
Nothing
4
Xothing
J
Notliing
4
Notliing
1
Nothing
4
Nothing
2
Nothing
2
Nothing
1
Nothing
4
ICpidermis
5
Nothing
2
Notliing
*' 2
Nothing
3
Nothing
1
Nothing
1
Nothing
2
Nothing
1
1
Kpidermis
Kpidermis
1
" . 1
I'phlermis
l';pi(k-rmis, Zygnenia, Mougcotia
2
l',|>idcrmis
ICpidcrmis, Spirogyra. Zygnenia. \'au-
cheria, Mougeotia, Navicula, Diatonia
S
Epidermis ?
5
Kpidermis
1
Kpidermis
5
Nothing
5
Kpidermis, Spirogyra, Oscillatoria. Dia-
toma, Synedra
5
Epidermis, e^g of Daphia, one of Simo-
cephalus, bit of plant tissue
" 3
Kpidermis
5
Kpidermis
1
Kpidermis
5
Epidermis
5
Epidermis
5
Ivpidermis
5
Epidermis
5
Epidermis. mud. Navicula, Kunotia,
^ Gomphonenia, Spirogyra
6
Kpidermis. Empididx ? 1
1
Epidermis
Epidermis
1
Epidermis
1
I^pidermis, egg of Simocephalus, 7 eggs
of Daphia, several statoblasts
5
Collembola 1
5
Tipulida: 1, Achene of Eleocharis acicu-
1
ICti'idermis, Oribatella 1
" 1
1
ICpidermis. plant fibers
1
Epidermis, Tassidx 1
1
Kpidermis
5
PhoridK 1, Chironomi(hc pupa. Coleop-
tera larva, Staphylinidx 1, achene of
Eleocharis acicularis
" S
I.ygwids 1, Attidffi 1
1
Jassids 1
1
Kpidermis, Crustacea egg, Carex seed
" 5
Nothing
" 5
Kpidermis
S
I'liidermis, Phoridae 1
6
Kpidermis. Lepidoptera larva. Diptera 2
Drosophilidx 1, Carabidae 1
46
Journal of Entomolog) and Zoolog)'
Table 4. Data for Rana palustris — Continued
95
28
96
28
97
28
98
38
<»9
28
FOOD
Kpidermis, Hydrachnidx 1, Jassidx 1,
Diptera 1, Colcoptcra I, few algal
filaments
Epidermis, Staphylinidx I
Lygn:idx 1, Xabidx nymph, Ephydri-
da; ? 1
Epidermis with Mougcotia, Haltica ? 1
Epidermis, Proctrotrupidx 1
Epidermis. Drassidx 2. Chalcididx 1,
Insccta 1
Diptera adult and larva, PoUcnia 1,
Khyncophora 1
Phoridx 1, Coleoptera larva
Epidermis
Uepidoptera larva
Diptera 1. Drosophilidx I. Carabidw 1
Planorbis 1, Heteroptera I, Staphylini-
dx 1
Epidermis, egg of Crustacea, Gamasidx
1, Mougeotia
Tipulidae 1, Dytiscidx 1, plant tissue,
Ulothrix pieces of chitin
Epidermis. Capsidx 1
Crustacea egg, Physopoda 1, CoUcmbola
I, I'ormicidx
Lymnxa 1, Laccobius 1. sand
Aphididx 1, Diptera 1, Rhyncophora 1,
winged seed
Epidermis, Collembola 1, Chrysomelidx
larva. Braconidx 1
Insecta lar\'a, seed
Insecta larva, Hydrophilidx 1
Epidermis. Heteroptera 1. Pollenia I,
Diptera 1
Epidermis, Lymnxa 1, Argiopoidca 2,
Jassidx I, Diptera 1, Drosophilidx 1,
1 Staphylinidx 1
Lymnxa 1, Capsidx 1. Psilopus 1, Lac-
cobius 1
Daphnia eggs, Liancalus 1. Carabidx I,
Haliplus 1. Juncus ovar»* with Lepi-
dop. larva. 2 Carcx achcnes
Insecta lar\'a, Cryllus 1. Jassidx I.
I Cercopidx I, Diptera adult and larva.
f Tipulidx 1. Acalyptcra 1
Epidermis. Tipulidx adult and larva.
Malipltdx 1, Proctrotrupid.T 1. small
leaf
Hclochara (Jassidx) 3, Tipulidx 1,
Colcoptcra I. Curculionidx l.Ephy-
(Irid.T 1. Chalcidid.T 1. Rraconidx 1
Argia nymph. Lyg.xidx 1, Rediviolus 1.
HcliKTh.-ira 2. Sphxrophoria 1, Lian-
calus 1, Colcoptcra larva, Philanthi-
dx 1, Rraconidx 1
Rann pi pirns SclireluT. The Meadow- or Li'«ip.Tril-fn>j;.
Total of 100 specimens. Lot 1, Isoetes Ponds, Chicago Bop, McLean, N. Y.,
July 22, 1916; lot 2, Bool's Backwater, Ithaca, Sept. \, 1912; lot 3, Taughannock
Pond. Ithaca, July 29. 1908; lot 4, Bool's Backwater. Ithaca, Aug. 18, 1906; lot 5,
Chicago Ponds. McLean. \. V.. July 30. I9|(i. Lot 1 collected by Dr. Wright and
myself; lot 2, by Hr. Wright; lot 3. bv Or. A. A. Allen; lot 4, by Or. Wright and
Dr. (;. IL Sabine; lot 5. by Dr. Wright and Dr. R. (i. (lilmorc.
Pomona College, Claremont, California
47
Table 5. Data for Rana pipiens
No. Body
Tail
Mouth
Ali. Can
Forele
gs Lo
1 29
44
tadpole
tadpole,
530 non
e 1
2 29
37
4U5
1
3 _7
37
210
4
4 27
36
360
1
5 28
48
changing
sti.niach.
150 pri.
sent 4
24 26 32
FOOD
Mud with Navicula, Diatoma, Synedra,
Spirogyra, Oedogonium, Ulothrix,
Vaucheria, Anursea, fibers, moss
Mud with Cladophora, Navicula, much
moss
Mud with Gomphonema, Cymbella,
Navicula, Cocconeis, Cyclotella, Dia-
toma, Synedra, Meridion, Spirogyra
Mud with Epithemia, Navicula, Acnan-
thidium, Vaucheria, pieces of leaves,
xylem spirals, broken tissue
Mud with Nitzchia, Synedra, Pinnularia,
Gomphonema, Cocconeis, Navicula,
Cymbella, Meridion, Diatoma, Cyclo-
tella, Closterium, Spirogyra, Oscilla-
toria, Cypridopsis, plant tissue
Diatoma, Navicula, Scenedesmus, Mou-
geotia
Nothing
Mud with Navicula, Ulothrix, fibers
Nothing
Nothing
Nothing
Nothing
Mud with Closteriuni, Pleurococcus, fil-
aments, plant tissue
Nothing
Nothing
Hpidermis
Nothing
Nothing
Nothing
Nothing
Epidermis ?, Oscillatoria. Zygneina, Spi-
rogyra, Cladophora, Synedra, Desmi-
dium, Gomphonema, Epithemia, Melo-
sira, mandible of insect larva, shell
of bivalve Crustacea
Nothing
Epidermis
Epidermis, Zygnema, Synedra
Epidermis
Nothing
Nothing
Epidermis
El)idcrmis,
geotia
Ilipidcrmis
Epidermis
Epidermis
Epidermis
Epidermis
Epidermis
Epidermis
Epidermis,
Epidermis
Nothing
Epidermis
Epidermis
Epidermis
Ivpidermis
Epidermis
J'jiHlermil
ICni.le
Chlor
Zygiiema, Ulothi
)ps (Oscinid.x) 1. Mil
1 . Heteroptera nymph, Jassida
1. Aphidids 2
Epidermis
Epidermis
Epidermis
j^pidermis, Drosophilidx 1
Epidermis
Epidermis, egg of Daphnia
Epidermis
48
Journal ot Entomology and Zoology
Table 5. Data for Rana pipiens — Continued
Xo. llody Tail
lipulirmis
hpidcrmis,
Ivpidermis
i'.pidcrmis
JCpidcrmis
I'.pidcnnis
I'.pidirniis.
Epidermis.
Upidcrmis.
Kpid.rmis.
lipi.lcrmis.
dx 1
Kddl)
Simuliidx 2
Dytiscidx 3 lar\'x
Dvtiscidx larva
Oribatidx 3, Collcmlx>la 1
Oribatidi 1
l.yinna:a palustris 1, Apliidi-
Orilwlid:
J
Oiptc
Mv
1. Collcmbola 1,
osophilida; 1
da* 2 nymplis. Apliidida- 2
Ltopliilid:i; 2, IMu.rtica 1. C
I.
)us 1
nplii
Kjiidcrmis. TItysanoptcra 1. Collcmbola
1, Aphididi 1, Dolicbopodida.- 1. Si-
muliidx 2, Braconidar 1
Nothing
Hpidemiis
Braconid.t 1
Kpidermis. Staphvlinida; 1, Collcmbola 1
I ymna^a 3. Muscidx 1
Epidermis. Collcmbola 1, Lymnjca 1
Epidermis. Tipulidx 1, Dytiscid^c 1
OribatitL-ie 3, Daphnia CKgs, Thysonap-
tcra 2, Collcmbola 3, Jassid.T 1, Psyl-
lid.T 1. Formicidx I
I'pidcrmis. \crmcs 1
I IclcritptiTa 1, llonioplcra 1. Capsidx I,
Muscid.x' I. Colcoptcra 4
Coleoplcr.1 ailult and larva, Capsida* 1,
Cercopidac 1
Hotcroptcra 1, Diptera I, Colcoptcra 2
Aranoida I. Capsid.T 1. Jassidw 1. Mus-
cida* 2. Colcoptcra 2, Hymenoptcra 1
Colcoptcra 2. Carabid.x 1. Formicida- 1
niptera I. Colcoptcra I. Clcridx I. lly-
nu-noptcra 1
Collcmbola I. Lampyridx 1. Stapltylini-
dx 1. Hymcnoptera
Piptcra 1. Colcoptcra 1. Chrysomclidx
1, Hymcnoptera 1, Philanthid.x 1
Doryphora 1. Clcridx 5, unidentifiable 1
Diptera arUiU and larva, Colcoptcra
larva. l>vtiscidx 1, Rhyncophora 2,
llraconidx 1
Jassidx 1, Carabidx 1. Chrysomclidx 1,
Staptiylinidx
Diptera 1. Colcoptcra 1. Muscidx 1
Oribatidx 1. Tassidx 1. Diptera adult
and larva. Carabidx 2. Stapliylinidx
I. Formicidx 1. Rraconidx
I.vmnxa 1. Mvcctopbilidx i. Phoridx 2
Diptera I. Colcoptcra 2 adults and I
Chrysomclidx 2, Proclrotrupi-
<lx 1
ICpidcrm^. I.vmn-a 2. Thvsonantcra 3.
Colkinhola I. Capsidx 1. Diptera !.
Dr<.s.M»bili<Ix 1. Colcoptcra I, Ily-
rlrophiliiJ.T 2. Chrysomclidx 1
I'jt'dirmis. snail. Carabidx 1. Drosophi-
HI,. 1. Formicidx 1
I iv. Ilctcroptcra I, Diptera 1.
.1. Rhvncophora I
P*.ytlidx 2. Diptera 1. Colc-
planldown
l*.r
.icnnidx 1
yllidx 1. Diptera 1. Carabidx I.
conid.T I, 4 pebbles
Pomona College, Claremont, California 49
Uyla cntiiji'r W'ied. The Spring Peeper.
Total of 32 specimens. Lot 1, Slaughter House Ponds, Ithaca, June 28, 1911;
lot 2, Cross-roads Pond, Ithaca, June 22, 1907; lot 3, Slaughter House Ponds, June
27, 1911; lot 4, Chicago Bog, McLean, N. Y., July 22, 1916; lot 5, pond on shore of
Lake Ontario, North Fair Haven, New York, July 30, 1916. Lots 1, 2, 3, by Dr.
Wright; lot 4, by Dr. Wright and myself; lot 5, by myself.
Table 6. Data for Hyla crucifier
\o. Body Tail Mouth Ali. Can. Forelegs Lot FOOD
1 11 18 tadpole tadpole, 66 none 1 Mud with Epithemia, Meridi
jilionenia, Diatoma, Eunoti:
Zygnema, Mougcotia
Ulothrix, Microspora, Crustacea eggs,
plant tissue, ParamcEcium
2 13 17 " " SO " 2 Mud with Zygnema, Oscillatoria, Micro-
spora, Mougeotia, Scenedesmus, Navi
cula
i 10 16 " •■ 17 " 1 Nothing
4 11 16 changing stom.-icli. 10 present J Nothing
.1 II 16 small •• 11 " 1 Nothing
() 10 14 changing " 11" 2 Nothing
7 11 13 " •• U " 2 Nothing
5 11 15 " " 13 " 1 Nothing
9 10 14 " ■■ 13 " 1 Nothing
111 11 11 small ■■ IJ '■ 1 Epidermis
11 10 11 " •• 12 •■ 1 Nothing
IJ 13 10 " •• II ■■ I Nothing
13 11 10 " " 15 " 1 Nothing
14 11 9 " " II ■■ 1 Nothing
15 11 S " " 12 " 1 Nothing
16 11 7 " stom. „■ int.. 12 " 1 Nothing
I" U 7 " " " 12 " 1 Epidermis ?
IS 12 I. large " " 13 " 1 Nothing
19 11 ft " ■■ ■■ 11 " 1 Nothing
2(1 12 5 " ■■ " 13 " 3 Nothing
21 11 5 " " ■• 12 " I Nothing
22 12 5 " ■■ •• IS " 3 Epidermis -
2i 12 2 " ■• •• 14 " 3 riiptera 1
24 14 II " •■ '• 24 ■• 4 Cfrcopid,T 2. Ciialcidids 1, Tchncumoni-
ds 1
25 14 II ■' " •• 20 " 4 Ilomoptera 1, Tipulidx 1
26 14 II ■• ** " 22 '* 4 i^iptera 2, Coleoptera 1, Tchneumonidx
1
27 14 0 " " " 23 " 4 Diptera 2, Coleoptera 1, Ichneumonida;
1
28 M 0 •• •• •• 14 ■' 1 Epidermis
29 11 n •■ •• ■• 17 " 1 niptera 1
30 'ft 0 " ■■ •• 20 " 5 niptera 1, Capsids 1, Phoridae 1
3' 'ft n " •■ " 19 " 5 niptera 2, Proctrotrupida: 2
32 15 0 " ■■ ■• 25 " 5 Insecta 2, Hymenoptera 1
Uyla ■versuolor Le Conte. The Common Tree-toad.
Total of 23 specimens. Lot 1, Ithaca, July 20, 1911; lot 2, Ithaca, July 22, 1908;
lot 3, Lake Ontario, North Fair Haven; lot 4, Ithaca, July 22, 1907. Lots 1, 2, 4, by
Dr. Wright; lot 3, by myself.
Table 7. Data for Hyla versicolor
No. Body Tail Mouth Ali. Can. Forelegs Lot FOOD
1 20 42 tadpole tadpole, 170 none 1 Mud with Pleurotsnium, Cosmarium,
Desmidium, Pediastrum, Scenedesmus,
Navicula, Pinnularia, Epithseraia, An-
abxna
50 Journal of Entomology and Zoology
Tabic 7. Data for Hyla versicolor — Continued
I1...1V Tail M.iiith
FOOD
, Ij ■>■> .. •• 170 '• 4 Mud with Navicula, rinnularia. Sync-
dra. Pediastrum, Sccncdcsmus, Cos-
inarium, Oscillatoria
1 in 20 '•^" ' ^'"'' "■■''' 1'lcurot.Tniuni, Cosmarium,
Dcsmidium, Pcdiaslrum, Navicula,
rinnularia, Anaba;na
A 15 22 chanKinR Moinacli. 30 |,rc>unl .' Nothing
5 14 II small ■• 20 " 2 Nothing
6 18 5 large " 20 " 3 Epidermis
7 19 J " stom. & int., 24 " 3 Nothing
g 20 2 " " *' 20 ** 3 Epidermis
9 ^0 4 " " " 20 ■' 3 Epidermis
lU ii 0 •■ •• •' 21 " 3 Insecta I, Diptcra 1. Clerid.-c 1
II 5o 0 •' ■• " 28 •■ 3 Epidermis. Oribatid.T 1, IJiptcra I.
Formicidw 1
l". ->! 0 " " " 23 ■• 3 Oribatid.x' I, Diptcra 1
j5 52 0 " " " 28 " 3 Psyllida.' 2, Trichoptcra I. Diptcra I,
Colcoplera Hymenoptcra 1
,4 -.. 0 •• " " 30 •■ 3 Insecta 2, larva 1. Nabidx 2, Diptcra 1,
Ichncumonida: 1
15 iQ 0 " " " 25 " 3 Tingitid.-c 2. Jassid.-e I. rsyllid.T 3, Dip-
tcra 1. Hymenoptcra 1, Chalcidid.T I
Ij ■>2 0 " " " 30 " 3 Insecta 3. Cleridx 1
17 ^1 n " •• " 42 " 3 Tingitidw I, Hymenoptcra 1, Apidae 1,
Mvrmicida:
lg 21 0 " " " 40 " 3 Tingilid.-e 17, Psyllida; 1. Colcoplera 1,
Ilyinenoptera I, plant down
19 20 0 •■ '• " 23 •' I Epidermis ?
20 20 n •• •■ ■• 23 ■■ 1 Epidermis
21 20 0 " " " 25 " 1 Epidermis
22 20 0 " " " 18 " 1 Epidermis
23 19 0 • 19 " 1 Epidermis
Rujn amcriianiis Holbrook. The Common Toad.
Total of 40 specimens. Lot 1, Dr. Wriglit and Dr. Reed, Cross-roads Ponds,
Ithaca, July 4, 1907; lot 2, Dr. \Vri);lit, Bool's Backwater, Itliaca, June 29, 1911;
lot 3, same, JuK 4, 19o7.
Table 8. Data for Bufo amerieanus
.No. Bo.ly Tail Mouth Ali. Can. Porelcgc I.ot FOOD
I II 12 tadpole tadpole. 110 none 1 Mud with Zygncma, Oscillatoria. Navi-
changing stomach.
tadpole
changing
Iwothirds
small
unc-haU
cula, Pando
rina. cj:gs of Crustacea
1
Mud with
Oscillatoria, Microspora,
Pandorina. '.
Navicula, Pinnularia, eggs
of Crustacea
present 1
Mud
none 1
Mud with Pandorina. Navicula
Mud with N.V
•-icula and plant tissue
present 1
Nothing
Mud with Nai
,'icula and plant tissue
.Nothing
l-'pidcnnis ?
Nothing
Nnlhing.
Nothing
Nothing
Nothing
Nothing
Nothing
Nothing
•Nothing
Epidermis ?
^tud. nothing
Mud
identifiable
Nothing idenlifinblc
.Nothing tdent
ifiable
Pomona College, Claremont, California 51
Table 8. Data for Bufo americanus— -Continued
No. Body Tail Moutli Ali. Can. Korc LcRs Lot FOOD
24 1(1 2 " ■■ ■■ 12 ■• 3 N'othing identifiable
25 9 2 ■' ■• ■• I J '■ 3 Kpidermis
26 ID 1 ■• •■ •■ 13 " 3 Kpidermis
27 1(1 1 ■• ■• '■ 11 •' 3 N'othing identifiable
28 10 i ■• •• " 13 " 3 Kpidermis ?
29 II 5 ■• •• ■• 13 " 3 Mud with Navicnla, masses of Plenro-
coccus
30 ') 0 " ■■ •■ 13 " 2 Physopoda 1, Insecta 1
31 11 .'; ■• ■• ■• 13 " 3 rulmonata 1
32 10 0 ■• " " 16 " 3 Kpidermis, iptera larva
33 II n •• ■• ■■ 13 " 3 Collembola 2
34 II 0 ■• ■■ ■■ 12 " 3 XothinK identifiable
3.'; II 0 •• •• ■■12 " 3 .\othinK
36 10 ■• •• ■• •• 10 •• 3 l';pidermis
37 10 0 ■■ ■■ ■■ 10 •' 3 lipidcrmis, Diptera 1
38 10 0 ■■ ■■ ■■ II •• 3 NothinK
39 9 n •■ •■ ■■ 10 •■ 3 N'othing
40 10 0 •• ■• •• 10 " 3 Kpidermis
Comparison of Tadpoles of the \^arioiis Species.
In the eight species used the tadpoles agree in being for the most part herbivorous.
The small mouth is provided with horny jaws and is used largely in nibbling off
Algae, bits of moss, and other plants, and in gathering up masses of ooze and mud
with the many diatoms and desmids to be found in such situations, and the occasional
Protozoa of the Difflugia and Arcella types.
Very often one sees statements such as made by Miss Dickerson that tadpoles,
especially of some species, are very "fond of any animal food available. Thus these
tadpoles act as scavengers and dispose of dead fish or dead tadpoles even, that would
otherwise become a menace to the living creatures of the pond." These statements
might indeed be made by almost anyone who has observed tadpoles to any extent. I
remember when a boy of reading that a good way of cleaning a skeleton of a small
animal like a mouse was to place it in a pond containing many tadpoles and it would
soon be nicely freed from the flesh. Experiment showed this to be more or less true;
but although I have studied many tadpoles in the series of forms now being discussed,
and although these come from many different ponds, the fact that in no case was such
animal matter found, leads me to believe that it is not so important a source of
food to the tadpole as is commonly believed.
Since all the tadpoles of the various species are aquatic and therefore in rather
uniform conditions, one would not expect their food to vary as much as does that of
the transformed individuals. The alimentary canal is invariably very long, in keeping
with the herbivorous habits; but almost entirely undifferentiated, no stomach nor
large intestine being evident. As long as the tadpole mouth is present the alimentary
Ganal is almost always filled with ooze and silt, a great part of which is inorganic.
Since the size of the mouth varies considerably with the species, one would expect it
to allow of more variation in food-habit than does any other one factor. I was par-
ticularly interested, therefore, to see what the largest animal form taken would be
and in which species it would be found. Unfortunately I did not have a very good
series of specimens with the tadpole mouth in the large bull-frog and green-frog, but
those examined showed almost no variation from the smaller species. One green-frog
did have a small crustacean (Ceriodaphnial), a meadow-frog contained a rotifer
(Anurfea), another had a crustacean (Cyt>ridupsu), and a peeper was found with
52
Journal of Entomology and Zoolog>'
many winter eggs of Crustacea. Aside from tlie few cases of Eitglena, Paramarclum,
Difflui/ia and Arcrlla met with, almost all of the remaining food was plant. No
attempt was made to make any quantitative observations on the plant materials
found. In number of individuals and actual amount of substance the diatoms were
very important; many desmids, some filamentous algae, and quite large amounts of
wood-fibers and tracheids, bits of leaves and other broken down plant tissues were
found. This is another bit of evidence in the rather vast amount which has now
accumulated to show the great importance of the diatoms in aquatic biology and
ecology. Table 9 shows in a relative way the frequency of occurrence of the various
forms of food.
TABLE 9
The Frequency Willi Which the Various Species of Tadpoles Contained
the Various Food-forms.
S[»ccinii:ns opened
UI.\TOMS
Epilhcmia
Navicula
Pinnularia
Diatoma
Syncdra
Xitzchia
Cymbella
Mcridion
Kunolia
(•nmphpncma
Miscellaneous
i-ii..\mi;ntous algae
/ygncma
lHothrix
Spirogyra
CjnHophora
MiMiKcotia
MKifllaneoils
I'.l.li; C.RKEN AI.GAE
< i-icillatoria
frog frog frog frog frog I'ccpcr toad Toad
IIKR Af.CAE
Closlerinm
Cosmarium
Prdiafltrum
Dcsmidiiim
riciirotxnium 1 .
Scencdcsmus
Mr
opcil
II.AC.KI.I.ATA
CKOTOZOA
Unril-KKA
CKI STACEA
IK.r.S Ol' CRUSTACEA
The figures given in this table indicate the number of stomachs in which the
various forms occurred, as no attempt was made to keep a count of the number of
times any one form was found in a given stomach.
Comparison of Young Transformed Individuals of the Various Species.
Jnsi a glance given at the data of the transformed individuals of the eight
*prcies as presented in the preceding pages, will show in a general way that their
fiMid consists largely of insects with some spiders, mites, and other forms, largely as
Pomona College, Claremont, California 53
has been reported for the adults by previous workers. I think it is worth while,
however, to go into more detail and to see, for example, whether the young frogs and
toads change at once to the more or less terrestrial habits of the adults or whether
they feed largely on the aquatic forms at first. Let us see, too, whether they are
limited very much by their size as to their range of food, and whether they begin
their predaceous habits at once or still feed on the diatoms and algae on which they
grew.
Perhaps a table comparing the different species will show most readily what we
desire. In Table 10 the animal forms contained in the stomachs examined have been
listed, the attempt being made to separate those which are without question aquatic
from those probably not taken in water. The CoUembola, young Anura, and insect
eggs might have been taken on water or not and are classed as doubtful. It is possible,
of course, that any of the winged insects might have fallen into water and have
been seized as they were struggling or floating on the water, but this could scarcely
have been true of many. Anyone who has watched transforming Anura knows that
they hop briskly about in the neighborhood of the pond and have every opportunity
to catch their prey in the air, from the surface of the mud, or from plants.
TABLE 10
Relative Numbers of Aquatic and Non-aquatic Animal Forms Found in
Transformed Individuals.
Protozoa
Water Snails
Crustacea and eggs 16
Water Mites 1
Odonata Nymphs 3
-Aquatic Hemiptera
Chironomid LarvK
.Aquatic Coleoptera 6
Rana Tadpole 1
TOT.AL AQUATIC FORMS 31
CoUembola 19
Eggs 2
Young Anura 2
Doubtful Forms 23
Wrmes
Land Snails
Land Crustacea 1
Mvriapods
Spiders 1
Land Mites 1"
* Land Insects "_
Adult Odonata 5
Thrips
Crickets
Land Hemiptera ^^
Scorpion Flies 1
Psocids 1
Lepidopterous Larvie 1
niptera ^
Dipterous Larvi
Coleoptera 21
Coleopterous Larvx
Hvmcnoptcra 6
TOT.SL XON-AOUATIC FORMS 71
TOT.M. AXIM.VL forms 125
Per Cent .\quatic Forms 25%
Per Cent Doubtful Forms 18%
Per Cent Land Forms 57%
Number of Stomachs 29
•Not furtber identifiable.
Bull- GrL
frog frog frog frog frog Peeper toad Toad
200
96
139
113
25
66
5
219
126
167
173
25
66
7
8%
21%
11%
13%
0%
0%
0%,
■/.%
2%
6%
22%
0%
0%,
29%
91%.
^6%
83%
65%
100%
100%
71%
54 Journal of Entomology and Zoology-
II is unfortunate that no more transformed individuals were available for the
toad, the per cents recorded for if are probably not worth a great deal; however, the
fact that no aquatic forms were found even in the five individuals studied is sug-
geslive and made understandable by the fact that young toads soon leave the ponds
by hundreds and at transformation time can be seen traveling toward the higher
ground in all directions. The absence of aquatic or even doubtful forms in both
species of llytn can probably be substantiated by the examination of larger numbers;
for young tree-frogs and peepers climb on plants above the ponds in which their larval
life was spent and, sitting on the leaves and branches of Iris, of shrubbery, or what-
ever is available, are ready to catch insects that crawl over the plants or come flying
to them.
It is noticeable, too, that the distribution of the forms eaten through many families
and orders is not nearly so great for these smaller species as for the species of Rana.
Dr. Wright* has shown that for the Ilhacan Anura the average lengths at transforma-
tion are as follows:
Ilufo americanus 9.6 mm.
Ilyta versicolor 16.0 "
llyla trucifer 1 1.0 "
Rana fiipiens 24.0 "
Rana patustrts 24.0 "
Rana sylvatica 16.0 "
Rana tlamilans 32.0 "
Rana mlfshfiana 53.0 "
TTie smaller size of some species naturally limits their food somewhat. The habit
in both species of llyla of sitting on plants, and their failure to hop about over the
ground as do some of the other forms may also have much to do with the explanation
of their eating fewer kinds of insects and other invertebrates such as spiders and
sow-bugs.
In the genus Rana a general tendency toward the habits of the adults is to be
observed; although the green-frog is a marked exception. One would expect young
bull-frogs to eat a rather large per cent of aquatic forms and the rather low per cents
given in I'able 2 for the wood-frog and meadow-frog are not surprising. But the
remarkably low per cent for the green-frog was hardly to be looked for. In this con-
nection a comparison with the data given by Surface" for the adult forms may be of
interest. His report lists the stomach-contents of Z') bull-frogs, of 107 green-frogs, 28
wood-frogs, 88 pickerel-frogs, 51 meadow -frogs, 17 peepers, 22 tree-toads, and 52
toads. By making a rough estimate of the forms which he lists 1 find that ihe com-
parison with the newly transformed is as follows:
'Wright, A. H.. I9H, I.ifc-hislorics of the Aniir.i of Iiliaci, New York. Cirncgie Institu-
tion of Washington.
'Surface. II. A., 1913. ICconomic features of .Amphibians of Pcnnsvlvnnin. Zoological Bull.
Pa.. Dcpt. of Agriculture, i;67\i2.
Bull-
frog
frog
Wood-
frog
Pickerel-
frog
Meadow-
frog
25 32
18 S
57 63
8 6
H 4
91 90
13 2
22 0
65 98
21 4
2 1
76 95
11 7
6 0
83 93
Pomona College, Claremont, California 55
TABLE n
Percentage of Aquatic Forms Found in the Food of Adults as Compared with
Newly Transformed.
adow-
Peeper toad
Aquatic
nmibtful
i-aquatic 57 63 91 90 65 98 76 95 83 93 100 100 100 100
In this table the figures express per cents, the one given first is for the young, the
second being for adult. It will be at once apparent that the bull-frog is by far the
most aquatic in feeding-habit, that the green-frog, although a form remaining close
to the water, lives verj' largely on non-aquatic insects, that the peeper, tree-toad,
and toad apparently eat practically no aquatic forms from the time that they
transform, and that the wood-frog, pickerel-frog, and meadow-frog leave the water
more gradually and always do have a small percentage of their food aquatic,
although not so much of it is so in the adults as in the young. Of all these
species the green-frog is perhaps the most surprising. Drake's'" results for the
meadow-frog, based on the most exhaustive study yet made and showing a total of
931 animals found in 209 stomachs, give about five per cent as being unquestionably
aquatic, so that his work agrees very well with the results given above.
Economic Bearing.
The economic application of a piece of work of this sort should be two-fold. As
new information is obtained regarding the food-habits of frogs, especially at trans-
formation, their life-history and propagation can be better understood. If frogs are
unable to eat at transformation, a fact which I think I have quite thoroughly estab-
lished, it is useless to feed them at that time. The second point of application that
comes to mind is that a study of the food of the newly transformed may show some-
thing as to the usefulness of the species in destroying harmful insects, sow-bugs, slugs,
and other forms. My data are hardly full enough nor important enough to go into
tliis in detail, but a more extended investigation of the food of the adults is worth
while from this standpoint. The results of other workers, such as Kirkland, Surface,
and Drake, do show that a great many harmful forms are destroyed. For more detail
their writings should be consulted.
Conclusions and Summary.
Eight species of Anura were studied during their transformation to learn some-
thing of their food-habits as larvae, as transforming individuals, and as young frogs
or toads. The species studied were as follows:
Rana catesbeiana Shaw. The Bull-frog.
Rana clamiians Latreille. The Green-frog.
Rana sylvalica Le Conte. The Wood-frog.
Rana palttstris Le Conte. The Pickerel-frog.
Rana pipiens Schreber. The Leopard- or Meadow-frog.
Hyla crucifer Wied. The Peeper.
Hyla versicolor Le Conte. The Tree-toad.
Bufo americanus Holbrook. The Common Toad.
"Drake, C. J., 1914. The food of Rana pipiens Shreber. Ohio Naturalist. 14:257-269.
56 Journal of Entomolopy and Zoology
In each species studied the same general tendencies are evident: ((1) The larval
alimentary canal is very long, but slightly differentiated in its various portions, and
filled with ooze and mud scraped up from objects in the pond and containing many
forms of diatoms, blue-green and green algte of filamentous and non-filamentous
types, small pieces of plant tissue, and bits of fiber and other slowly decaying material
to be found in ooze. Very few tadpoles were found with any animal food, the excep-
tions having a few small Crustacea, Protozoa and Rotifera.
(2) After both pairs of legs are evident and the horny plates of the tadpole
mouth are shed, the tail is found to be gradually absorbed and the alimentary canal
decreases to about one-tenth of its larval length at the same time that it widens
anteriorly to form the stomach and posteriorly to form the large intestine. During
this transformation period the mouth increases to about six or seven times its former
size and there is practically no feeding done. The epidermis is apparently shed
rather frequently as the tail is being absorbed; for its presence in the alimentary canal
during the final stages of transformation is so frequent as to be quite universal in the
larger species and occurs in all those studied.
(3) Ailer these changes have been just about completed the young frog or toad
begins life as a carnivor, apparently taking anything movable yet small enough for it
to handle. Occasional bits of plant-down and small feathers testify to the attractive-
ness of a moving object. Almost all groups of invertebrates and some vertebrates are
represented in the diet, the largest per cent being insects, crustaceans, spiders, sow-
bugs, and snails. Some individuals do contain pieces of plant tissue, sand, mud, and
other inactive objects, but these seem to be accidental, often occurring where ground
beetles or similar forms have been eaten.
(4) The newly transformed individuals show a decided tendency toward the
habits of the adults; the toad, tree-toad, and peeper eating almost nothing of an
aquatic nature; the meadow-frog, pickerel-frog, and wood-frog eating some aquatic
forms, a few per cent more than do the adults of their species; of the other two species,
both of which are considered i|uilc n(|tjalic in habit, the green-frog lias about nine-
Icnlhs of its food non-aquatic and tlic bull-frog about three-fourths non-aquatic.
Uy way of summary, then, the tadpoles of the species of .^nura studied for this
paper are largely herbivorous, the transforming individuals do almost no feeding,
and the young frogs or toads are mostly carnivorous. These changes in habit are
made possible by great changes in the alimentary canal and mouth.
The Central Nervous System of Three
Bivalves
WIl.I.IAM A. IllI.TON
Lima Deliscens.
The central nervous system forms a rather compact mass of nervous tissue, with
certain special local thickenings where nerve cells are abundant. As in Pectin, as
described by Drew, the visceral ganglion is the largest, but it is not so widely sepa-
rated from the other ganglia as in Pectin. Neither is it so complicated in structure.
There are, on each side, three main branches from the visceral ganglion, the most
caudal goes over the adductor muscle to the mantle. The next, the smallest main
branch, goes to the gills, Vfhile the last branch, the largest, is chiefly a mantle branch,
which divides after leaving the ganglion.
The cerebro-pleural ganglia are connected medio-caudally by a looped com-
missure, the other large medial branch on each side runs to the rather large pedal
ganglion, while near it is the small otocystic branch, much as in Pectin. The large,
more cephalic branch runs towards the mouth region and gives off a number of
branches, about seven.
The pedal ganglion is made up of two nearly distinct parts and from each of
these lateral parts a branch runs into the foot.
The visceral ganglion is more complex than the others in structure, but there
are only a few distinct fiber tracts.
In all tlie ganglia, the cells are large or ganglionic and small or ordinary nerve
cells.
Sunset Clam, Psammohia (■alijitrnica
The cerebral ganglia are of fair size and not widely separated. There is a
cephalic branch supplying the mouth region and palps and a more ventral branch
also on each side, supplies neighboring parts. The commissure between the two
ganglia is rather narrow considering the size of these centers.
The Pedal ganglion is small and gives little indication of being divided into
two parts. The two connectives come to it and two rather large branches leave.
The visceral ganglion is large and especially well developed. This is because
of the large siphons and their necessary abundant nerve supply. The siphons are
capable of being extended some distance from the shell. The ganglion is complexly
lobed on superficial view. There are on each half at least six little lobes which
represent to some degree groups of nerve cells. On each side in addition to the
large connective branch there are branches as follows: (1) a large branch to the
gills, (2) a large trunk which divides again into mantle branches. One of its branches
going to the dorsal siphon, (3) a small dorsal branch, (4) a small ventral branch, (5)
another large mantle branch which sends some strands to the ventral siphon, (6)
another large mantle branch, (7) a small branch to the posterior adductor muscle.
58
Journal of Entoniolop,' and Zfxilogy
'^^://;;!|
■VV4J. ,
FJR. I. Crnrral Kanglia of Lima X9.
FiK- 2. Section of Orrhral ganglion X70.
Fig. 3. Section of Pedal ganglion X70.
Fig. 4. Chief ganglia of Piddock X9.
Pomona College, Claremont, California 59
Sections were made of the ganglia. The cerebral ganglia were found to be more
complex than those of some other bivalves. This was shown in the differences in cell
groups and greater complexity of the central fiber masses. The individual cells differ
greatly in size, but they average somewhat larger than in some other bivalve forms
studied.
The pedal ganglion, although not so complex, also shows differences between cells.
There are large multipolar cells and among these are small ones of various sizes.
Tlie processes of the larger cells may be traced into the fibrous mass for some distance.
The visceral ganglion is composed of two large lateral parts closely fused.
There are numerous commissural bands binding the two sides, but the chief fusion
is by more or less individual fibers. Cells inclose the whole ganglion and as in the
other centers they are of large and small size. The cell areas of the larger cells are
mostly localized on the dorsal and upper surfaces, but the lower end of the ganglion
has some large cells. The large cells are especially found in the neighborhood of the
larger branches, those branches supplying the mantle and siphons and it seems that
some of the larger cells are concerned with supplying these characteristic parts.
The California Piddock, Parapholas californica Conr.
The ganglia were dissected in medium sized individuals. The cerebral ganglia
are about as in other bivalves. The ganglia are quite widely separated. Besides the
commissures connecting them and connectives to lower ganglia there are several
branches to the mouth region from the upper and lateral sides.
The visceral ganglion forms a larger mass than any other of the ganglia. There
is verv little indication of right and left halves. Closely joined to it is the small pedal
ganglion.
Microscopic examination of serial sections bring out further details.
The cerebral ganglion is simple in structure. There are a large number of cells
in proportion to the fibres in the center of the ganglia. As in many other molluscs,
there are many small cells and a few much larger ones, but these last are not abun-
dant. In the large cells it is not difficult to determine fine fibrils and strands from the
smaller cells near by. There is also a very complex mingling of strands from the
central fibrous mass. Some of the fibers are small, some are larger. The appearance
of these larger cells is much as described by .'\pathy. The cells in the ganglion are
chiefly multipolar.
The visceral ganglion is the largest and most complicated. Caudally it sends
two thick nerves backwards. These are its chief branches for a long distance; they
do not branch. The two sides of the ganglion are joined by many cross fibers and
there a few bundles in distinct commissures. Most of the cells are small, but there
are a few of the larger type. The cells form a rather uniform sheath all about the
ganglion, but here and there we find special cell areas. The fibers are much less
evenly disposed and present a very complex mat in every part.
The large cells in some cases have a symmetrical distribution. There are certain
individual lateral cells of this sort, also some dorso-central ones which seem to occupy
60
Journal of Entomolop,- and Zoolog\'
^^:si?^
Fig. II. Cerebral ganglia. X9. Sunset clam.
Fig. 12. Pedal ganglion. X9. Sunset clam.
Fig. 13. Visceral ganglion from tlie side. (a) ("onncctivc branch, (b) dorsal
hrancli, (c) gill branch, (d) branch to posterior adductor muscle, (e) mantle branch,
(f) matitlc and ventral siphon branch, tg) small mantle branch ?, (h) mantle and
dorsal siphon branch. Sunset clam.
Figs. 14 and 15. Othtr views of the visceral ganglion, lettering as in Fig. 3. X9.
Pomona College, Claremont, California
61
,.*
\\
'%
> )
Fig. 16. Cerebral ganglion, section. The commissure shows. X70. Sunset clam.
Fig. 17. Longitudinal section of a pedal ganglion. X70. Sunset clam.
Figs. 18, 19, 20, 21. Various sections of visceral ganglion. The dorsal side is
up. X70. Sunset clam.
62 Journal of Entoniolog\ and Zoology
charactcriMic positions. Also on the dorsal side llicre are (wo peculiar fiber masses in
symmcirical positions.
The pedal ganglion is small and just in front of the visceral. Il is almost a
pan of the visceral and closely applied to it. It has two chief nerves on the cephalic
side. Its central fibrous mass is slight.
Explanation of Figures.
Fig. 4. Chief ganglia of Piddock. The cerebral ganglia are above, the vis-
ceral mass with the small pedal ganglion attached below. X9.
Fig. 5. One cerebral ganglion with part of the connective. X350.
Fig. 6. One cerebral ganglion, a branch above, the connective to the left, the
commissure branch to the right. X70.
Figs. 7, 8 and 9. Sections through three levels of the visceral ganglion. The
dorsal side is up. X70.
Fig. 10. Section through the pedal ganglion with the two connectives as isolated
pieces each side. The dorsal side is up. X70.
VOLUME TWELVE NUMBER THREE
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
SEPTEMBER, 1920
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT 0/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
A List and Some Notes on the Lizhards and Snakes Represented
IN THE Pomona College Museum — Raymond B. Coivles - - 63
The Central Nervous System of an Unknown Species of
Marine Le^ch — William A. Hilton 67
Central Nervous System of a Centipeip— /^r/Aar S. Campbell - - 69
Microscopic Studies of the Water of the Claremont-Laguna
Region — Geneveive Corwin - . - 72
Preliminary List of Microscopic Life in Fresh Water Pools
Around Lacuna Beach - - 74
Preliminary List of Microscopic Life in Fresh Water Around
Claremont 76
General Reactions of a Centipede — Susie Case ----- 79
Notes on the Central Nervous System of a Free-Living Marine
Nematode — William A. Hilton - - - 82
Entered Claremont. Cal..Post-OfBce Out. 1, 1610, as second-class matter, under Act of Congress of
March s. 187» /
\^ NOV 1-7 1939
Journal of Entomology and Zoology
EDITED BY POMONA COLLEGE, DEPARTMENT OF ZOOLOGY
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and entomological journals, proceedings, transactions, reports
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The pages of the journal are especially open to western ento-
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Authors of articles longer than a thousand words will receive
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Address all communications to
The Journal of Entomology and Zoology
William A. HUton, Editor
Claremont, California, U. S. A.
A List and Some Notes on the Lizards
and Snakes Represented in the
Pomona College Museum
Raymond B. Cowles
The purpose of this article is to give a general idea as to the distribution of
snakes and lizards from the desert regions of Southern California, with a few observa-
tions on their habits. It is also an enumeration of the snakes and lizards which may
be met with in the region about Claremont.
The list has been compiled from specimens in the Pomona College Museum only,
and the writer is well aware that not all the specimens from the Claremont and
desert regions are represented. No effort is made to give the limits of the range of
the specimens nor to give any conclusions as final. In those cases where a list is
given of the places from which specimens were taken, it is merely to show that
the range is at least of that extent.
Testudo agassizi (Cooper).
One of these desert tortoise was taken at Ludlow, California, towards the last of
April, 1920. It was found out in the open at the base of an alluvial fan, and made
no effort to escape capture. It is being kept alive with a view to study its habits so
far as possible under artificial conditions.
Dipsnsaiirus diirsalis (Baird and Girard).
Taken from fifteen miles east of Blythe Junction, .'\pril 2, 1920, in the sand hills.
A second specimen was taken 45 miles west of Blvthe, in a sand wash, on April 4,
1920.
The main habitat of this lizard seems to be the sand hills or sandy country, and
it takes refuge in the holes of rats when menaced.
During August of 1919 they were seen in pairs and seemed to be breeding.
Observations seemed to show that a given pair occupied the same territory and
rarely traveled far from it. They were seen most on the hottest days, feeding on
the leaves of some of the low desert shrubs. Upon being frightned they would drop
from the branches and run rapidly, with the entire body raised from the ground, to the
nearest burrow, where they would remain for half an hour or more before reappearing.
On cloudy days, even though the temperature remained above 100° F. they were
seldom seen and appeared to be very sluggish, sometimes allowing one to approach
to within a few feet of them before running.
Their food seemed to be almost exclusively plants, and they preferred the leaves
of an alfalfa plant which happened to be growing near their chosen range. During
an entire summer, June 25 until September 25, they were seen eating insects only
once. The specimen eating the insect escaped and it is not know what insect it might
be, though from a distance it appeared to be one of the ."^crididae.
64 Journal of Entomology and Zoology
I'ma nolala (Baird).
Only one specimen of this beautiful lizard is found in the itiuseum, and i( was
taken in the sand hills 15 miles east of Blythe Junction, April 2, 1920. The lizard
is very shy, running rapidly to the shelter of a burrow in the sand, at the least threat
of danger. (This seems to be between I', nolala and U. scoparia.)
Calhaurus I'fntralis ventralis (HallowelU.
This lizard appears to be one of the most numerous and widely distributed of
the Colorado and Mojave deserts, having been found in almost every type of country
with the exception of the rocky hills and mountains, from Victorville to Needles and
south to the Mexican Border in Imperial Valley. In the Providence Mountains they
were found at an altitude of over a thousand feet.
In the Imperial Valley they were found to burrow, or push down into the sand
at the approach of night. Here they remained until sunrise of the next day. At the
approach of danger they jump from the sand with such suddenness as to give the
impression of a small explosion.
The distribution as given above is not intended as a limit to their range but
merely a note on their presence in those places.
Crolaphylus coUaris hailfyi (Stejneger).
This lizard is represented by three specimens in the college collection. One taken
from near the Bonanza King Mine, Providence Mountains, March 31, 1920; another
from the N. E. spur of the Turtle Mountains, and a second and smaller one from
the same place, April 1, 1920.
These lizards were found on the rocky hill-sides and were very active and rather
shy. Their strong jaws and great speed fit them for the predaceous life which they
lead. In the largest specimen was found an eight inch CncmiJoplionis tigris ligris,
partially digested.
Crolaphylus v:izlizrnii (Baird and Girard).
Two specimens were taken at the grass fields between Blythe and Mecca, on
April 2, 1920.
These specimens were found skulking under the branches of the creosote bushes.
They are very rapid runners, and are predaceous. Their coloring blends admirably
into the mottled shade where they lie in wait for their prey. A ten-inch Cnemidophorus
ligris tigris was taken from an eleven inch specimen. Their biting ability was well
proved upon the collector who picked up one of the specimens which had been only
wounded. One bite tore through the skin of the first finger, causing a decided flow
of blood.
Sauromatits aUr (Pumeril).
One specimen taken in the lava rocks east of Ludlow, March 30, 1920. Two
specimens taken among the rocks in the N. E. spur of the Turtle Mountains.
These lizards, which are not fast runners, are usually found near some crevice
in the rocks in which they take refuge upon the approach of danger.
The two specimens taken in the Turtle Mountains, April 1, 1920, were found as
a pair, and when first seen appeared to be in copula. This gives some suggestion as
to the lime of breeding.
Pomona College, Claremont, California 65
Via Stansburiana clegans (Yarrow).
Several specimens were taken during the first week in April, and they seem to be
farily common throughout a large part of the Mojave and Colorado deserts, in Cali-
fornia at least.
Sceloporus magisler (Hallowell).
One specimen taken 35 miles east of Mecca, California, April 2, 1920. Other
specimens taken during July and August, east of Holtville, California. These lizards
seem, to prefer the brushy country or the neighborhood of trees, into which they
climb when frightened. The specimen taken east of Mecca was found on the ground
beneath a cactus.
Phyrnosoma plalyrhinos (Girard).
Representatives from five miles west of Amboy and Needles, California. Without
an exception they were found on the dry gravelly washes or in the sand not far from
washes.
Xantusia vigilis (Baird).
Three specimens from east of Victorville, and one from the Providence Moun-
tains, near Bonanza King Mine, March 30, 1920. These specimens were all found
beneath the bark of prostrate yuccas.
Cnemidophorus tigris tigris (Baird and Girard).
These lizards appear to be one of the most common found on the Colorado and
Mojave deserts in California. Their rsnge is extremely varied, specimens being
taken from, and between, Victorville, Needles, Blythe, the Mexican border in Imperial
Valley, and Palm Canyon. These localities are not given as the limits of the range
but places within the range from which we have specimens. Specimens were taken
in the Salton Sink 265 feet below sea level, and from the Providence Mountains at an
approximate altitude of 2,800 feet above sea level.
Sonora occipitalis (Hallowell).
One specimen taken at the grass-fields, between Blythe and Mecca, California.
When taken it was traveling out in the open and in the heat of the noon sun, April
3, 1920. It was found on a gravel wash and when approached it struck in all
directions, though apparently it did not open its mouth upon striking the hand. It
appeared to be blinded by the sun and unable to tell from which direction it was
menaced.
Bascanion jlagelliim frenatum.
Two specimens, both taken near Mecca, Imperial Valley, April 4, 1920. Both
these specimens were somewhat lighter than specimens taken from the region around
Claremont, California.
One of these snakes was obtained under rather unusual circumstances, which
incidentally involved the collecting of a Cnemidophorus tigris tigris. The lizard was
shot but not killed by the collector, and while watching for an opportunity to kill
the lizard without the use of a second shot, the snake was seen gliding in the same
direction as the lizard, and suddenly attacked and seized it, when both were added
to the collection.
66 Journal of Entomology and Zoology
Crolalus mildifUi (Cope).
This specimen was collected by Dr. Hilton ami Dr. Miinz of Pomona t'ollepc, at
Forest Home, San Bernardino Mountains, June ", 1919.
Crolalus ifraslfs (Hallowell).
One specimen taken at Needles, California, April 1, 1920. These snakes seem to
be almost rniircly restricted to the sandy areas of the desert, rarely wandering from
them, and then only for a short distance, its mode of locomotion admirably fits it for
the tvpe of country which it inhabits. The ordinary snake finds difficulty in rapid
motion over the loose and shifting sand, since part of the tractive power comes from
a bracing of each loop of the body against that part of the ground which is posterior
to the loop, and through the movement of the central portion of the body against
the surface of the ground. It can readily be seen that a shifting and loose surface
would seriously hinder the progress of the ordinary snake. The "Side-winder,"
Crolalus teraiirs, instead of progressing as do ordinary snakes, longitudinally, pro-
gresses laterally, leaving separate tracks, each paralleling the other, and angling in the
direction in which the snake is moving. Each track is approximately the length of the
snake making it, and is wavy, that is, a series of "S" shaped loops. The tracks
give no sign of any part of the body moving from one mark to the other, which gives
the impression that the snake jumps the 3 to 6 inch interval between the tracks. Such is
not the case, however. When the snake is moving, the body is kept partially looped
and the advance seems to he through the advancing of the head and tail, while the
rest of the body is rested on the intervening loop, supporting the rest of the body,
the weight then seems to be shifted to the head and tail and the rest of the body
advanced, the whole progression being a series of graceful and contiiuiors movements.
This sterns to be the mode of progression.
Crolalus alrnx (Baird and Ciirard).
Taken at Mecca, California, April 4, 1920. Foiuid in ilic arrow w;.'ed where
it seemed to be fairly common.
In addition to the above list of specimens from the desert region there remain
that from the vicinity of Claremont, California, which is as follows: i'la slanshuriana
htsfteris, Richardson; Sielufiorus oiiiJentalis hi-sfrialus, Hallowell; I'/irynosoma
blainvillii hlainvillii, Ciray; Gerrlionolus scinc'uauda txfhhii, Baird; .Innitlla fulclira
puldira, (tray; .Innirlla (iluchra nigra, Fisher( doubtful location. Specimen not
labeled. Another from Laguna Beach August 1, 1920) ; Cnrmidopliorus ligris slej-
iiff/rri. Van llenburgh; I'lrsliodon skillonianum. Baird and Girard; l.iclianura rose-
ojusca. Cope (two taken from vicinity of Claremont and one from east of Victorville
by \V. M. Pierce); Tliamnnfiliis nnlino'iiles hamonJii. Kennicott; OiaJnf'liis amahitis,
Baird and (Jirard; l.nmfrnpfltis pyrnmrlana inultiiincia. Yarrow; l.amprnprllis
lioylii, Baird and (iirard; R/iinoi hrihis Irconlei, Baird and Ciirard; llypsiglena
orlirorhyndius. Cope; Salvadora liexalefiis, Cope (taken in Imperial Valley 10 miles
cast of Holtville) ; Coluhrr eonslrirlor vfluslus, Baird and Girard; Coluhfr flagellum
frrnalus, Slejneger; Coluhrr lalrralis, Hallowell; P'lluopliis tatrniffr ralrniffr. Blain-
vllle; Crolalus orrganus, Holbrook.
The Central Nervous System of an Un-
known Species of Marine Leach
Wll.l.IAM A. HILTON
Tlie little animals from vvliicli this study was made were obtained during the
summer of 1920 at Laguna Beach. Two times when a number of Mysis shrimps were
brought in with towings these worms were found attached by the posterior sucker to
the side of the crustacean. At first it was not clear to which group of animals these
small creatures belonged. It was not until a number of the specimens had been cut in
series that their nature was learned. Externally they seemed unsegmented, although
the body had many circular rings when contracted by reagents, but these rings were
evidently not marks of segmentation. Internally at first there also seemed to be little
trace of metamerism, but when the nervous system was examined a clearly defined
chain of ganglia was evident.
The mouth is at the base of the large anterior sucker, and it is back of this that
the ganglia may be seen. The chief ganglion is the suboesophageal composed of about
four parts fused and closely applied to the next ganglion below. The brain or supra-
oesophageal ganglion is unimportant; in fact, it is the smallest of all. There are sixteen
sinmple ganglia forming the ventral chain back of the suboesophageal and the seven-
teenth ganglion or last of the chain. The last center, or the seventeenth, is made up
of at least three simple ganglia fused and is the second most important center. It
supplies the structures of the large posterior sucker.
Some of the points of special interest in the nervous system of this creature are;
1. Lack of true metamerism except in the nervous system.
2. The large number of simple clearly defined nerve centers. About four centers
are represented in the suboesophageal, sixteen separate ganglia and at least three
separate centers for the last ganglion. In all then there are at last twenty-three centers
in the nervous system.
3. The small size of the supraoesophageal ganglion or brain.
4. The large size of the suboesophageal ganglion and the last ganglion.
5. No special sense organs were located.
The specimens were from 4-8 mm. in length and, although small, were sexually
mature. The identity of the species will be considered at another time.
(Contribution from the Zoological Laboratory of Pomona College.)
v> ^ — ^
'ii^\t<i%iLi^\.
"%«s
\4k0^^^
EXPLANATION OF FIGURES
Below, the general position of llie ganglia is shown. On the left above is an
enlarged longitudinal section of the upper ganglia and just below it a cross-section
through the brain and sulxrsophageal with the oesophagus in the space between. The
two upper central figures are longitudinal and cross-sections of about the tenth
ganglion. The last figure to the left is a longitudinal section of the last ganglion.
The dorsal side is up in all the figures. The sections are all enlarged 170 times, the
figure of the whole animal is enlarged 20 times.
Central Nervous System of a Centiped
ARTHUR S. CAMPBELL
The central nervous system of S. Polymnrfi/in Woods, is especially studied in the
present paper.
Hymonds (1898) considers the development giving especial note to the homologies
of this system. Newport (1843) gives some notes in regard to the brain. Saint-Remy
(1890) gives considerable detail especially in regard to the finer structure of the
brain of S. Morsitans L. Case (1920) has shown something of the behavior of
S. Polymop/ia and indirectly the arrangement of never tracts.
Ordinary dissections and the occaasional use of a binocular microscope proved
the most useful.
Successful stains were Heidenheim's and Delafield's Haemetoxlins. HgCl.- or
AgN'Oa seemed the best fixers. Tracheae were studied without reagents immediately
after exposure.
In S. Polymorpha the supraoesophagal ganglion or brain comprises three paired,
fused divisions or lobes. Large branches extend from the antenna] lobes into the
antennae. The ocular lobe fuses with this and is distinctly larger and less markedly
hilohate. This lobe sends out nerves to the oceili. The labro-fontal division is under-
neath the ocular lobe and entirely fused with it. It innervates the labrum.
The supraoesophageal ganglion in S. Polymorpha is large. It is anteriorally
connected with the brain by two circumoral connectives. Ten principal, paried
nerves are connected with this ganglion. The anterior pair extend into the mandibles.
The second pair supplies the first maxillae, the third runs to the second maxillae. The
fourth pair innervates the maxillipeds. The fifth pair supplies the prehensorial
feet.
The remaining somites are supplied by simple, similar ganglia, equally spaced
but well separated by connectives. The third and fourth ganglia area almost fused,
due to the foreshortened segments in which they are located. There is no histological
difference between them and other abdominal ganglia. One ganglion only is present
in each somite. Altogether in S. Polymorpha there are twenty-four ganglia.
Each abdominal ganglion gives off eight nerves. There is no ventral nerve. The
first pair of branches supplies the tergular muscles, the second the walking legs, the
third the sternal muscles and the fourth supplies the spiracles and tracheae.
The two caudal ganglia present special interest. Four principal branches run
from the first of these. The first supplies the tergal muscles, the second the sternal
muscles while the fourth supplies the anal legs. Additionally, two preanal connectives
join with a small ganglia about half the normal size of the others. Four nerves
extend from this last small ganglion into the sphincter and other anal muscles.
In general the superficial tracheal distribution is rather definite and much
resembles that of the insects. The brain is rather poorly supplied by but two main
tracheae on either side which break up into a number of tracheoles which run into
the antennae and optic lobes. In contrast to this, the suboesophageal ganglion is
supplied dorsally by three tracheaae on each side.
70 Journal of Ent<iinology and Zoology
The abdominal ganglia are eacli supplied by two ventral tracheae. The dorsal
tracheae send vessels throughout the length of the branches on the dorsum of the
ganglion. Each ganglion is well supplied by numerous small iracheoles.
The two candal ganglia present seme ditTcrences in the distribution of tracheal
eleme.^ls. The dorsal surfaces of the twenty-third and twenty-fourth ganglia is sup-
plied by six tracheae. X'eiitrally there is cne principal branch suppling both by
numerous tracheoles.
Histologically the brain and other ganglia resemble mich those of the more
generalized Insects. I have found little difference in my specimens and those figured
by Saint-Remy (1890) of 5. Morsilans. The cellular masses of all my preparations
seem much less than those figured by Saint-Remy. The fiberous area of the brain
contains some indication of lobular masses. There are at least two sizes of cells
noticeable.
In the abdominal ganglia the fiberous mass occupies rather more than half the
bulk. The cellular area, composed of several sizes of cells, is closely crowded.
The caudal ganglia contain less bulk of the fiberous mass and a large area
of cells. The cells here seem to be all of approximately the same size and type.
In all preparations, the nuclei appear large, the nucleoli show prominently.
Tigroid substances was noticed in a few of the larger, better stained cells, especially
in the brain. Fibrils were seen to enter into certain cells, and touch the nuclei.
CONCLUSIONS
1. The central nervous system of S. Folymorp/ia is ccmpcscd of twenly-four
generalized ganglia. The brain is less comple.x than that of the insects.
2. Of the three primitive elements of the brai.i two only are externally apparent.
3. Tracheae supplying the central nervous system art definitely arranged.
4. The functional cells of the central nervous system are of several sizes, the
fiberous mass makes up the greater bulk of the ganglion. The cellular area is external
and relatively less abundant.
5. Nuclei are large, nucleoli arc well markeil. Fibrils appear to come into
contact with nuclei.
BIBLK)t;R.\PHY
Casf, S.: General Reactions of a Centipede. 1920
Journal of F.nlomology and Zoology.
Ilrymon.ls: 1S9S
Zur Kntwicklungsgcschichte der Chilopoden.
Silzungslicr K. press. .Akad. Wissenschaft.
Die Enlwicklungsgeschiechte der Scolopender.
Zoologia, Heft. 33.
\fiifiort, George:
Structure, Relations and ncvelupmcnl of the Nervous a id Circulatory
Systems - - - in Myriopoda and Macrourous .'Xrachnida.
Philos. Transactions of the Royal Society.
Saint-Remy, G:
Contributions i I'ctude du ccrveau chez les Arlhropoiles Trachcates.
Archiv. Zool. Exper. Tome V. suppl. 1887-90.
(Contribution from the Zoological Laboratory of Pomona College.)
1901
Fig. 1. Brain and suboesophageal ganglion; tracliea? black. X6.
Fig. 2. Twenty-third and fourth ganglion. .\5.
Fig. 3. Abdominal ganglion. X5.
Microscopic Studies of the Water of the
Claremont-Laguna Region
CESEVEIVE CORWIN
The climatic conditions in Southern California where these studies were made,
are unusual in that the rainy season occurs during the winter and early spring and
there is practically no rainfall for the rest of the year. About 10 to 15 inches is the
average yearly amount. V\'ith this small amount of precipitation, most of the
streams dry up completely and the permanent pools diminish in size. This fact has
a profound effect upon the life contained in the water. Just how this effect works
out has not been determined. Some forms are able to dry up and still retain life,
while others are killed by lack of moisture. Almost all the studies recorded in this
paper were made on permanent pools and streams.
Studies of the microscopic life of the Claremont-Laguna region were made in the
early spring and summer, those of the Claremont region in February, March and
April; and of the Laguna region during the last half of June and the month of
July of the previous year.
Considering the two places as a whole, in general there were more green algae
than blue-green; more algae than Protozoa, the amoeboid Protozoa being fewest in
number; the flagellate a little more numerous and the ciliate most frequent, both in
species and individuals. The rotifers were rather rare, but were quite varied in
form, from the simply constructed, active Cnlurus to the beautifully ciliated fixed
Floseularia. The Castrolricha were very rare.
The chief difference between the Caremont and the Laguna regions is the abun-
dance of aquatic life. This might be caused by the fact that most of the pools
studied around the Laguna were close to the shore and the water may have been
brackish. As a rule they were more stagnant than the Claremont water, with the
exception of the Laguna Lakes. Perhaps the seasonal change may have had some-
thing to do with this difference. The Claremont studies were made over a period
of time twice as long as the other and much earlier in the season. However this
may be, in almost every group there were more species in the Claremont region
than the Laguna and in all other cases there were at least as many, with the
one exception of the one desmid found in Claremont and not in Laguna. To
summarize the comparison: There were twice as many species of algae in the Clare-
ment region as the Laguna; the same number of blue-green for both localitis but
four limes as many green in Claremenl. The diatoms were quite numerous and
varied in form in both places but there were only half as many species in Laguna.
As mentioned before, one desmid was found in Clarement and none in Laguna.
The Protozoa were quite abundant in both regions, there being three times as
many in Claremont as in Laguna. In Claremont the amoeboid were twice as
numerous as at Laguna. There was a larger proportion of beautiful complicated
forms in the Claremont region. There were three species of Sitnlor in Claremont
and only two in Laguna. The restless little Euplotn, the graceful Spiroslomium, the
beautiful Slylonye/iia are illustrations of the variety of ciliates in Claremont.
Pomona College, Claremont, California 73
There were one-half more rotifers in Claremont than Laguna. However, Laguna
had in comparative abundance the very interesting form, Rotifer neptunis. This form
is quite long and slender when extended, with two rosettes of cilia and a quite
unmistakable Neptune's trident at the end of the tail. It is very collapsable, telescoping
down to one-third of its extended length. This was peculiar to the smaller Laguna
Lake.
Claremont showed several specimens of Brarh'wnus. I am not certain of the
species but the name must stand for want of a better one. It was a large form with
two magnificent wheels of cilia and two short slender arms, each bearing a tuft of
cila. When the animal drew in the wheels of cilia at least one of these arms
remained exposed. It was rather sedentary, fastening its two small toes to a piece
of algae and bending its flexible, stout body in different directions to search for food.
Only one Gastrotricha was found in the Claremont region while this same genus
(Chaetonolus) was found in two different places and more than one individual
was seen.
Microscopic Crustacea were rather rare, only one (Cyclops) being found in the
Sulphur Spring at Laguna. Three other kinds were found in the Claremont region,
two in the South Hills, the other at Puddingstone Canyon and in the Puente Hills.
One water mite was found in Claremont in a temporary pool and in no other
place.
The comparison between the temporary and permanent pools is not adequate on
account of the scarcity of data. In a general way, there is a smaller variety and
number of forms in the temporary than in the permanent pools. Streams and perma-
nent pools are similar in the amount of life they contain.
Preliminary List of Microscopic Life in
Fresh Water Pools Around
Latruna Beach
I. Algae
7.
.•\mphora
A. Blue-green
.■\lgae Pool
1.
C)»cillatnria : found in
8.
Cymbella
Algae Pool
Sulphur Spring
Smallest Laguna Lake
Smallest Laguna
LaKe
Largest Laguna Lake
Largest Laguna
Lake
2.
Spirulina
9.
Pinnularia
Smallest Laguna Lake
Smallest Laguna
Lake
Largest Laguna Lake
Salt Spring
Algae Pool
10.
CJomphonema
Laguna Canyon Pool
Smallest Laguna
Lake
3.
Nostoc
U.
Closterium
Smallest Laguna Pool
Smallest Laguna
Lake
Algae Pool
12.
Pleurosigma
Laguna Canyon Pool
Smallest Laguna
Lake
4.
Nodularia
13.
Epitliemia
Laguna Slough
Smallest Laguna
Lake
5.
Pliormidium
Smallest Laguna Lake
in. Protozoa
.'\. .Amoeboid
B. c;
1.
ireen
Cladophora
1.
Amoeba
Algae Pool
Salt Spring
2.
Nuclearia
2.
Synedra
Salt Spring
Laguna Canyon
B. Flagellate
Smallest Laguna Lake
1.
Luglena spirogyra
3.
Ankislrodesmus
Smallest Laguna
Lake
Algae Pool
2.
Euglena sp.
Laguna Slough
Laguna Canyon
4.
Spirogyra
Laguna Slough
Laguna Canyon Pool
3.
Phacus longicaudis
Laguna Slough
Smallest Laguna
Lake
Smaller Laguna Lake
Laguna Canyon
5.
Sccnedesmus
C. C
iliate
Largest Laguna Lake
I.
CJonium
6.
Navicula
Laguna Canyon
I.
Smallest Laguna
Flexiphyllum
Lake
Algae Pool
Smallest Laguna
Lake
Salt Spring
3.
Condylostoma
Sulphur Spring
Smallest Laguna
Lake
Smallest L. Lake
Largest Laguna [
Lake
Pomona College, Claremont, California 75
Laguna Canyon \'I. Rotatoria
4. Paramoecium A. Rotifer neptunis
Laguna Slough Smallest Laguna Lake
Algae Pool B. Rotifer citrinus
Sulphur Spring Sulphur Spring
Salt Spring Salt Spring
5. Lacrymaria Laguna Canyon
Laguna Canyon C. Diplois
6. Stentor (fixed) Smallest Laguna Lake
Laguna Canyon Stentor jj. Colurus grallator
(moving) Smaller L. Smallest Laguna Lake
Lake Salt Spring
7. Vorticella Laguna Canyon
Smallest Laguna Lake Algae Pool
Laguna Slough r n- .• j •
" " L. .Notius quadricornus
Algae Fool cm t i i
" Smallest Laguna Lake
Laguna Canyon
Volvox
Largest Laguna Lake
F. Philodina roseola
Laguna Canyon
Gastrotricha
Flat Worms
1. Jensenia *'^' Chaetonotus
Laguna Canyon , Laguna Canyon
Smallest Laguna Lake
Round Worms
Smallest Laguna Lake ^'"'- Copepoda
Laguna Canyon A. Cyclops
Algae Pool Sulphur Spring
Preliminary List of Microscopic Life in
Fresh Water Around Claremont
The numbers after ilie genera refer
made.
I. Algae
A. Blue-green
2. Oscillaloria 3, 6, 7, 9.
3. Nostoc 1, 9.
4. Merismopedia 2, 3, 4.
5. Spirulina 2.
6. Masiigonema 3.
B. Green
1. Vaucheria 11.
2. Cladophora 4, 8, 11, 13, 14, 15.
3. Clamydomonas 2, 3, 7, 8, 13,
14, 15.
4. Gonium 13, 14, 15.
5. Spirogvra 1, 3, 5, 7, 8, 9, 14.
6. riothrix 13.
7. Mougeotia 13.
8. Mydrodlctyon 2.
9. Pediaslrum 2, 12.
10. Scenedesmus 2.
11. Chlorospliaera 3.
12. Chactophora 8, 9, 14.
13. Zygnema 7, S.
14. Chlorngonium 7.
15. Myxonema 7.
C. Diatoms
1. Navicula 1, 2, 3, 4, 7, 8, 9,
11, 12. 13, 14, 15.
2. Epithemia 8, 9, 12, 14, 15.
3. Synedra 1, 2, 3, 4, 5, 7, 8, 9,
13, 14, 15.
4. Cocconela 1, 3, 5, 8, 9, 13,
14, 15.
5. Siurella 7, 8, 12, 13, 14, 15.
6. Ciomphonema 1, 2, 3, 4, 5, 7,
8, 13, 15.
7. Amphora 2, 7, 8, 13, 15.
8. Nllzschia 1, 4, 7.
9. Rhoicosphenia 3, 4, 7.
10. Tahellaria 7.
11. Cymbclla 2, 13, 15.
to the stations where the collections were
12. Selenastrum 2.
13. Cyclotella 2, 4, 13.
14. Pinnularia 2.
15. Encyonema 3, 8, 13, 14.
16. Denticula 3, 5, 8, 11, 14.
17. Eunotia 4, 13, 14.
18. Plagiogramma 4.
20. Triceratium 4.
D. Desmids
1. Cosmarium 3.
2. Closlerium 2, 5, 8, 9, 13, 14.
15.
II. Protozoa
A. Amoeboid
1. Aclinosphaerium 3.
2. Amoeba limax I, 13, 14.
3. Amoeba 3.
4. Acanthocystis 13.
5. Noclearia 2, 3.
B. Flagellate
1. Euglena 5, 6, 7, 13, 14, 15.
2. Peranema 6, 7.
3. Notosolemus 6.
4. Eulreptia 6.
5. Atractonema 7.
6. Phacus 7, 15.
7. Astasia 3.
8. Ccphalothamiiium I, 13, 14,
15.
9. Irceolus 14.
10. Heteroncma 14.
11. Trentonia 15.
C. Ciliate
1. Vorticella I, 2, 5, 6, 8, 13. 14.
2. Stentor 13, 14, 15.
3. Stentor polymorphus 14, 15.
4. Linotus 14.
5. Colpodium 12.
6. Leucophrys 7, 14.
7. Euplotes 1, 3, 6, 8, 13, 14.
Pomona College, Claremont, California
77
8. Cinetochiliim 3, 4, 14.
9. Cotluirnia 14.
10. Paramoecium 1, 2, 3, 6, S, 15.
11. Pleuronema 2, 7, S.
12. Stylonychia (long) 2, 3, 7, 8,
12.
13. Stylonychia (oval S, 15.
14. Oxytricha 5, S, 10.
15. Chilodon 1, 3.
16. Chaenia 1, 2, 3, 6, 8, 11.
17. Atractonema I.
IS. Ophryglena 1.
19. Frontonia 1.
20. Glaucoma 2.
21. Condylostoma 3.
22. Coleps 3, S, 15.
23. Colpoda 8, 12.
24. Metopus 8.
25. Halteria 7.
26. Spirostomium 6.
27. Blepharisma 15.
28. Opercularia 15.
III. Rotifera
1. Pleiirotioclia 8.
2. Philodina 6, 8.
3. Gastropus 1.
4. Diplax 3.
5. Diplois 1, 13, 14, 15.
6. Brancliionus 5, 13, 14, 15.
7. Rattulus 14.
8. Floscularia 14.
9. Diascliiza 13, 15.
10. Melicerta 15.
IV. Gastrotriclia
1. Chaelonotus
V. Crustacea
A. Ostracoda
1. Cypris 7, 9.
2. Herpetocypris 11.
B. Cladocera
1. Alonella 11.
(Contribution from the Zoological Laboratory of Pomona College.)
This peculiar load was br()ii);lit iiilci the lahoralory by Mr. M. Wymaii. I'lie
drawing is by Mr. K. (."rosswliilc. The load lived for some liiiu' and a few things
were learned about its extra leg with llie f.vo feet.
1. It was capable of feeble movements of the leg and feet.
2. There was no true joint at the junction of the fifth lej; with the body.
3. The extra leg was draj^gcd alo.ig with no attempt made to use it in any way.
4. The extra leg could be used as a brace when the toad tried to climb from
a jar.
(C'ontribtitiiin from the Zoological I.ali.raiory of Pomona t'nllcge. )
General Reactions of a Centipede
SUSIE CASE
This paper deals with the locomotion and general reactions resulting from
experimentation upon the nervous system of centipedes. The nervous systems of these
forms are very good for such experimentation, as the ganglia are distinct and widely
separated.
There seem to be but three or four papers on the subject — two of these being on
the physiology of the brain and not behavior, and one, "On the Movements of Milli-
pedes and Centipedes" by E. Ray Lankester. I should like to mention several points
which were observed along this last line. The locomotion of the centipede can be
better emphasized by comparing it with that of the millipede. In the millipede one
of the most apparent characteristics is the movement of the legs in waves, the pairs
on opposite sides moving together, identically. The legs form groups of two pairs
to a segment and these start the motion from the tail end forward. From five to
eight distinct waves can be counted when all the legs are in motion. Millipedes
move straight forward. On the other hand, the centipede as stated by Lankester,
"contributes the serpentine stroke to the process of locomotion." It does not have
the distinct waves mentioned in locomotion of the millepede. The legs on the opposite
side do not move identically but are antogonistic in phase; and move in perfect
harmony unless there be some injury to the nervous system, which controls locomotion.
I agree with Lankester that it is most probable that the condition presented by the
centipede in locomotion is a higher development than that shown by the millipede.
The wave movement suggests a type found in lower invertebrates.
Th reverse locomotion of the centipede is very interesting. Most of them persist
in going forward and yet in testing to find some definite result, I have discovered
that occasionally they will, with persuasion, go backward. Most often, however, they
turn the entire body instead of reversing the movements of the legs. On the other
hand, all millipedes with persuasion will reverse for a short distance. \\Tien one
goes backwards, it reverses the motion of the waves also, causing them to go from
head to tail instead of from tail to head.
I have mentioned the two main observations of general behavior as to locomo-
tion and shall now go on to the definite experiments which were made on the
centipede to test specific reactions.
First as to the method: The specimen to be operated upon was pinned out on
cork — the pins not being put through the centipede but across in a sufficient number of
places to hold it firmly. The cut was made from the dorsal side into the nervous
system. We tried not to make the external cut any larger than was absolutely
necessary. When in doubt as to the position of the injury, we examined the animal
after death.
The experiments and results are as follows:
Experiment /...Twelfth connective cut on right side. Results:
1. Some lack of movement in legs near cut and on same side, probably due to
injury of muscles.
80 Journal of Plntomology and Zoology
2. Tests to see ivlieilier stimuli carried from tail end to head end on injured
side. Anal leg pinched. We have the suggestion in this that the impulse travels up
and crosses over to the opposite side at the injured point, causing the head to turn
to the right. On the uninjured side the impulse is able to travel up without crossing.
The reaction was <|uicker than on the injured side.
3. Acetic acid on antennae of injured side. Reaction on opposite side at anal
end first. Acetic acid on antennae of uninjured side. Reaction on same side at
anal end.
4. When stmulated below cut, both sides respond equally well. .\\\ of these
tests show that movement is deferred on the injured side.
Experimrnl II. Similar results obtained by cutting connective in fourteenth seg-
ment on right side.
ExprrimenI III. Cut two connectives of twelfth segment. Results:
1. Specimen was turned on its back. It could turn over above injury without
aid, was helpless back of injury.
2. Moved legs vigorously above injury; dragged others.
3. Antennae sensitive to touch, causing response back to injury.
Experiment II'. Results similar to experiment three obtained by cutting two con-
nectives between last two ganglia.
Experiment I'. Connectives cut between brain and sub-ganglion. Results:
1. Stimulated antennae. No response.
2. Stimulate anal leg. Impulse traveled along slowly, causing all legs to move.
This seems to be a muscular reaction rather than one controlled by the nervous system.
3. One response in which I was very much interested was that the centipede,
as a result of this particular experiment, reversed movement with apparent ease.
Experiment I' I. Two alternating connectives cut. Results:
1. Specimen very active. Tests showed good crossing of sensation paths.
Experiment I'll. Four cuts alternating excepting for second cut. Between cuts
one and two connectives not severed on either side. Results:
1. Test to see whether stimuli carried to brain. Very slight stimulus at anal
leg, caused only reaction in legs back of injury. Strong stimulus, caused stimulus
to go to brain but it was very slow, due to the number of injuries. The stimulus
had to cross at several points.
2. There is apparent separation of brain from anal end by injuries. The legs
in front of injuries in constant motion, while those in back are quiet.
3. Stimulated head region. Result is a very active reaction, which takes place
almost immediately, back to the injured part. There was much delay here. CJradually
the response extended farther down.
Experiment I'lll. C<mneclive cut on left side in tifth segment from head. Con-
nective cut on right side in fourth segment from tail. In this experiment I wanted
to test for time of response when cuts are on opposite sides and quite a distance
apart. Results:
1. Anal legs stimulated. On the right side it took longer for the response at the
head end. On the left side it was carried immediately to brain. This was probably
due to the position of the segment where crossing over took place.
Pomona College, Clareinont, California 81
2. Legs stimulated at center of body. Anal end drew up on tlie side stimulated.
This reaction toolf longer on the right side, because the stimulus had to cross at the
injury.
3. From the injury of the nervous system of the muscles, the specimen moved
with a swinging motion. It could reverse its movements.
Experiment IX. About one-third of the brain was removed, the right connective
was severed between the brain and the next ganglia, all connections with the eye
were severed on the same side. Results:
1. No co-ordination of leg movement. Legs interfered with one another.
2. At first, no sense of correct position. As willing to stay on back as normal
position.
3. Most noticeable result was that it reversed movement with apparently as
much ease as it went forward. It traveled the length of the dish. This centipede
lived twenty-four hours.
Experiment X. Removed sub and supra ganglia. Results:
1. Had better co-ordination of leg movement than one with one-third of brain
removed (Experiment IX), however, it needed stimulation for movement. A slight
jar of the dish was stimulus enough for the reaction. After this experiment the
centipede lived sixty hours, thus showing the injury to be less of a shock than in
experiment nine.
Experiment XI. The centipede was cut into nearly equal parts. This last
experiment is of a different type but results are along the same line as others.
Results:
1. In tail half there seems to be co-ordinated reaction of legs, suggesting that
the symmetry has not been interfered with. It turns toward side stimulated. Tail
end remained alive a little over two hours.
2. The head end was again cut into two parts. The central section was active
and remained alive for two hours. The head end was very active. It had initiative
to move without being stimulated, which power the other two parts did not have.
The head end remained alive three hours.
GENERAL CONCLUSIONS
1. The head ganglia seem to be necessary to initiate movements.
2. The body ganglia are rather independent centers for local control, and
complete co-ordination is possible without the head.
3. The stimuli travel up and down the nervous system, both on the side
stimulated and on the opposite side.
4. In case a connective is served on one side, the stimulus is capable of crossing
over to the other side but the reaction is somewhat delayed.
5. When alternate connectives are severed for some distance, the stmulus,
although delayed, passes from one end to the other. The delay is increased according
to the number of connectives severed.
6. Centipedes as compared with millipedes do not as a rule reverse the
movements of the legs, but unilateral injuries to the brain seem to permit the reverse
movements upon stimulation.
(Contribution from the Zoological Laboratory of Pomona College.)
Notes on the Central Nervous System of
a Free-Living Marine Nematode
WILLIAM A. HILTON
The species studied was the one which Is most abundant at Laguna Beach among
Algi and in sand at low tide. It corresponds closely to Enoplus brevis Duj.
The nervous system has several features not described in related forms. There is
a concentration of the central nervous system. There is a single large ganglion or
brain in the snout above the mouth, from this two connectives pass ventrally to join
the broad ventral nerve band in the mid-ventral line, while the only other longitudinal
nerve noted was the very small mid-dorsal. Lateral nerves were not found.
The head or snout ganglion is provided with three eye spots, and unpaired dorso-
median and a pair of latero-ventral ones. The sensitive region is so placed as to
receive stimuli from above by the median eye and from below by the lateral eyes.
The eyes are little more than concave pigment spots imbedded in the mass of the
ganglion. .X number of fibers pass from the ganglion forward to supply the thick
sensory epithelium of the tip of the snout.
The ganglion is rather complex in structure. It has a central and somewhat
ventral mass of fibers surrounded on all sides by nerve cells and fibers mingled. There
are two centers composed each of cell areas surrounding a fibrous mass; these seem to
be associated with fibers connected with the sensory epithelium of the snout and they
resemble slightly the olfactory areas of certain Invertebrate brains.
The dorsal nerve trunk Is not cellular. The ventral nerve trunk is thick and
broad. Ventrally it is nearly fused with the underlying cells of the body-wall, while
dorsally It is bounded by a closely applied muscular layer. The nervous tissue itself
is traversed by heavy lines which in part may be merely supportive in function, the
lighter strands, both transverse and longitudinal, are branches from the rather abun-
dant cells which are for the most part located ventrally.
(Conlrihulion from the '/.nnlogical Lahoratory of Pomona C.nllrgf.)
(FIG. 1.) EXPLANATION OF FIGURES
The figure above is a reconstruction of the head end of Enoplus, showing the
position of the nervous system. The lower figure at the left is of a section through the
whole body of the worm, showing the dorsal and ventral nerve bands. Both these
figures enlarged 75 times. The drawing at the right is from a section through the
head ganglion, enlarged 170 times. The dorsal side is up in all the figures.
(FIG. :. . EXPI-ANATION OF FIGIRES
The figure abo\-e is chrough the snout and ganglion of Enoplus. The central
figure is a drawing of a cross-section of the ventral nerve band. The loivest figure is
from a longitudinal section of the ventral nerve band with the muscular layer above
and the body-wall below.
The dor^al side i> up in all the fieures and all are enlarged 275 times.
VOLUME TWELVE NUMBER FOUR
>*
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
DECEMBER, 1920
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT of ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
New Species of Crane-Flies from the United States and
Canada — Charles P. Alexander , - - 85
Notes on Pacific Coast Pycnogonids — If. A. Hilton - - - 93
UcA MusiCA— /. CaUhi-ell. If. Dimmt 94
Lepidopia Myops — J. Ciildiccll 95
Eremita Analoga — Ha-tiard Lorheer 96
The Nervous System and Sense Organs. I. II and III — Ifillinm
A. Hilton ---------- - - - 1 to 14
Entered Claremont, Cal.. Post -Office Oct. 1. 181V. as second-class matter, under Act of Congress of
March 9. ItlTS
W NOvi? 193B
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Journal of Entomology and Zoology
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The Journal of H^ntomologv and Zoology
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ClaremoDt, California, U. S. A.
New Species of Crane-Flies from the
United States and Canada
(Tipulidie, Diptera).
By Charles P. Alexander, State Natural History Survey, Urbana, Illinois.
Most of the new species described in the present paper were found among material
sent to the writer for identification. I am greatly indebted to Mr. W. L. McAfee and
to Mr. F. R. Cole for the privilege of studying and describing many of the species
included in this paper. Two interesting forms were collected in southern Illinois
during the season of 1919 by Mr. Malloch and the writer.
Family Tipulidae.
Subfamily LimnobiinsE.
Genus Dicranomyia Stephens.
Dicranomyia terrie-nov,e sp. n.
General coloration gray, the praescutum with three dark brown stripes; antennse
dark brown throughout, the flagellar segments short-oval; wings with a heavy dark
brown pattern, including five large costal blotches; Sc short, basal deflection of Cul
far before the fork of M.
Male. — Length about 5.5 mm.; wing, 7.6 mm.
Female. — Length about 7.5 mm.; wing, 7.7 mm.
Rostrum dark brown; palpi brownish black. Antennae dark brown, the flagellar
segments short-oval, clothed with an abundant pale pubescence. Head bright silvery
on the front, duller on the posterior parts of the head; a conspicuous brown line on
the vertex.
Pronotum dark brown. Mesothorax very deep, the mesonotum gibbous. Mesonotal
prasscutum light gray with three conspicuous dark brown stripes, the broad median
stripe indistinctly split by a capillary line; scutum gray with the lobes dark brown;
scutellum and postnotum gray, the latter with a delicate brown median line. Pleura
light gray with an indistinct brownish longitudinal stripe extending backward from
the fore coxae; a similar line on the mesosternum. Halteres yellow, the knobs dark
brown. Legs with the coxae small, gray; trochanters dull yellow; femora brownish
yellow, the tips indistinctly darker; tibiae and tarsi brown. Wings whitish subhyaline
with a heavy brown and grayish pattern, as follows: five dark brown blotches along
the costal margin, the first near the wing-base, the third at the tip of Sc and the origin
of Rs, the fourth at the tip of Rl , the last at the tip of R2-\-3, suffusing the wing-apex;
the first three of these markings reach the costa and pass into cell R; the fourth
(stigmal) is rectangular, connected with a blotch at the fork of Rs ; narrow brown
seams along the cord and the outer end of cell 1st M2 ; large brownish gray clouds
along the margin at the ends of the veins and at the anal angle of the wings.
Venation: Sc short, ending just beyond the origin of Rs, Sc2 indistinct, apparently
somewhat removed from the tip of Scl, this distance about equal to the basal deflec-
86 Journal of Entomolog)- and Zoology
tion of Ml+2; basal deflection of Cul far before the fork of M, this distance about
equal to the basal deflection of Atl+2.
Abdomen dark brown, the posterior margins of the segments broadly silvery.
Ihihilal. — Newfoundland.
Holotype, 9, Spruce Brook, August 8-12, 1912 (G. H. Englehardt), (No. F3192).
Allotopotype, 3.
Paratopotype, 9-
Type in the collection of the American Museum of Natural History.
Dicranomyia lerra-novir differs conspicuously from all the described American
species of the genus. Its vicarious Palacarctic representative is D. decora (Staeger)
of Northern Europe. Superficially it bears a marked resemblance to Geranomyia
rostrata (Say), from which the structure of the mouth-parts and the slightly different
venation will separate it.
Genus Flliptera Schiner.
Eltiplera illini, sp. n.
General coloration brown, the pleura yellowish; cell Isl M2 open.
Female. — Length about 5 mm. ; wing, 6 mm.
Rostrum pale brown, the palpi dark brown. Antenna with the scapal segments
pale yellowish, the flagellum black; flagellar segments oval with a sparse white
pubescence and verticils that are a little shorter than the segments. Head dark
brownish black.
Thorax dull yellow, the thoracic dorsum with the stripes brown and entirely con-
fluent, shiny, only the lateral margins of the prxscutum yellowish. Halteres dark
brown, the base of the stem more yellowish. Legs with the coxas and trochanters dull
yellow; remainder of the legs brown, the base of the femora paler. Wings gray, the
stigma indistinct; veins dark brown. Venation: Sc rather short, ending about opposite
two-thirds the length of the long sector; Sc2 proximad of the origin of the sector, the
distance about equal to the basal deflection of Cul ; basal deflection of R-/+5 almost
square and in one wing of the type strongly spurred at the angle; cell Isl M2 open by
the atrophy of the outer deflection of Mi, Afl-\-2 before m about one-half that beyond
this cross-vein; basal deflection of Cul just before the fork of M.
Abdominal tergites dark brown, the sicrnites yellowish.
Habitat. — Illinois.
Holotype, 9, Makanda, Jackson County, June 4, 1919 (Alexander).
Type in the collection of the HIinois State Natural History Survey.
The unique type of F.lliplera illini was found in the "Ozark" region of Illinois
while Mr. Malloch and the writer were engaged in an entomological survey of this
section. The genus Elliptera was hitherto represented by two species from Europe
and two species from North America west of the Rockies. The occurrence of the
genus east of the Mississippi River was quite unexpected and breaks the hitherto dis-
continuous distribution of this curious genus of crane-flies. The present species differs
from its American relatives in the open cell Ijt M2, a character possessed by both of
the European forms.
Pomona College, Claremont, California 87
Genus Orimarga Osten Sacken.
Orimarga wetmorei sp. n.
General coloration black; thoracic pleura and lateral margin of the praescutum
striped with silvery; legs pale yellowish brown, the tips of the feinora a little paler;
wings subhyaline, the veins pale brown; tip of Rl atrophied or indistinct; deflection
of R4+5 very long.
Sex, female? — wing, about 4 mm.
The type is badly discolored. The general coloration is a dark brownish black;
basal segments of the antennae paler, the flagellar segments nearly globular.
The mesonotum has the extreme lateral margins of the praescutum narrowly
silvery, the pleura with a broad silvery longitudinal stripe, this type of coloration
being similar to that in 0. argenteopleura. Legs light yellowish brown, the tips of
the femora indistinctly paler; tarsi darker. Wings subhyaline, the veins pale brown,
more yellowish along the costal margin. Venation: Sc moderately long, ending at
about one-third the length of the long sector; Rs strongly arcuated at its origin; tip
of Rl atrophied or retreated back almost to the tip of Scl ; r very long and strongly
arcuated; basal deflection of R4-\-5 very long, strongly arcuated at its origin, more
than half the length of Rs ; cell MS deep; r-m far beyond r.
Abdomen dark brownish black, the apex broken.
Habitat. — Florida.
Holotype, Sex?, Paradise Key, February 22, 1919 (Alex Wetmore).
Type in the collection of the United States Biological Survey.
0. luetmorei is the sixth American species to be described, the second from the
United States. The fly differs conspicuously from O. arizonensis Coq. (Arizona) in
the coloration of the legs and body and in the venation. It is much more like O.
argenteopleura Alex. (Guatemala) which has the thorax similarly colored; this latter
species is considerably larger, with dark brown legs and a very distinct venation
(tip of Rl short, persistent; basal deflection of R4-\-5 short).
The species is dedicated to the collector, Alex Wetmore.
Genus Erioptera Meigen.
Erioptera (Erioptera) oregonensis, sp. n.
Size large (wing of the male over 7 mm.) ; general coloration brown, including
the halteres; wings with a strong brownish suffusion.
Male. — Length, 6 ram. ; wing, 7.3 mm.
Rostrum and palpi dark brown. Antenna; dark brown, moderately elongate,
clothed with a dense white pubescence, the verticils of the more terminal segments very
long. Head dark brown, more grayish brown around the eyes.
Mesonotum dark brown with indistinct stripes on the prasscutum, the lateral
margins of which are indistinctly paler; humeral angles not noticeably brightened;
tuberculate pits small, widely separated; scutum, scutellum and postnotum sparsely
yellowish gray pruinose. Pleura dark brownish black, gray pruinose. Halteres long
and slender, dark brown, only the base of the stem a little brightened. Legs with the
coxae dark, grayish pruinose; remainder of the legs dark brownish black, only the
trochanters and the bases of the femora a little brighter. Wings with a strong grayish
88 Journal of Entomology and Zoology'
brown suffusion; stigma dark brown; an indistinct brown cloud along r-m and the
deflection of R-/ + 5; veins dark brown. Venation as in the subgenus, the 2nd Anal
vein strongly sinuate.
Abdomen dark brownish black with a paler brown pollen. Hypopygium a little
brighter; pleurites short and stout, sparsely setigerous; two pleural appendages, the
outer appendage larger, the outer end flattened and enlarged, along the margin with
four parallel rows of fine comb-like points; inner appendage paddle-like, the blade
suddenly enlarged, provided with a few setigerous punctures, at the extreme tip with
an additional, powerful, curved bristle. Penis-guard straight, tapering gradually to
the blunt tip; gonapophyses with the apices produced laterad into conspicuous tri-
angular blades with the points directed laterad.
Habitat. — Oregon.
Holotype, S, Tillamook, March 26, 1919, (A. C. Burrill).
Genus Ormosia Rondani.
Ormosia subcornuta, sp. n.
Belongs to the meigrnii group; closely allied to O. cornula (Doane) but the veins
stouter, the stigma distinct, and the details of the male hypopygium very different.
Malf. — Length, about 3.5 — 3.8 mm.; wing, 4.3 — ^.7 mm.
Female. — Length, about 3.8 — ♦ mm.; wing, 5 mm.
Rostrum and palpi dark brown. Antennae moderately elongate, dark brovrnish
black, the scapal segments slightly paler brown. Head gray, provided with con-
spicuous yellow seta;.
Thoracic dorsum brownish gray without distinct stripes, the lateral margins more
yellowish; tuberculate pits shiny black, located close together, the distance between
them less than the diameter of one. Pleura brown with a strong gray pruinosity; a
large tuft of yellow setsc between the base of the wings and the base of the halteres
and a second group immediately ventrad of the halteres. Halteres yellow. Legs
with the coxae dark, gray pruinose; trochanters dull brown; remainder of the legs
dark brown, the bases of the femora a little brighter. Wings subhyaline; stigma
large, dark brown; veins stout, dark brown. Venation: cell 1st M2 open by the
atrophy of the outer deflection of Mi; 2nd .Inal vein slightly sinuous on its distal
half, converging toward the lit .Inal vein.
Abdomen dark brown. Male hypopygium with the pleurites stout, provided with
numerous conspicuous setigerous tubercles that bear long yellowish seta; which become
more elongate and stouter toward the tips of the pleurites; outer pleural appendage
subglobular, armed with from 4 to 8 powerful, acute spines, the terminal spine large,
along the outer face with microscopic, appressed denticles, the basal spine on the
inner side of the appendage largest, strongly incurved; inner pleural appendage
long, slender, with a strong spine before the tip to produce a bifid appearance. The
most lateral pair of gonapophyses are sinuous, with a group of two or three teeth or
spines on the inner face some distance before the tip, the slender apex bevond these
slightly curved; the proximal pair of gonapophyses are almost straight, very slender,
the lip with numerous indistinct denticles, at the extreme base with a few conspicuous
spines; an additional pair of gonapophyses whose apices are conspicuously flattened,
with the point of the blade directed laterad and slightly ccphalad. Ninth slernite
Pomona College, Claremont, California 89
with a broad spatulate blade, as in the mei/jrnii group of this genus, the apex deeply
notched medially.
Habitat. — O regon .
Holotype, $, Forest Grove, March 26, 1919, (F. R. Cole).
Allotopotype, 9.
Paratopotypes, 2 5 s; paratypes, 1 <5 , 1$, Hillsboro, April 1, 1919, (F. R. Cole).
This little species is evidently the Western representative of the common 0.
meigenii (O. S.) of the Eastern States, its general appearance being very like that
species. In the structure of the male hypopygium, however, it runs closes to O.
cornuia (Doane), which may be told by the different color of the wings and the
structure of the hypopygium.
Genus Gonomyia Meigen.
Gonomyia (Gonomyia) coloradica, sp. n.
Belong to the blanJa group, closest to mallwsoni Alex.; general coloration yel-
lowish, the praescutum with three broad, confluent stripes of reddish brown; wings
with the petiole of cell M2 long; male hypopygium with the structural details very
diflFerent from those in G. mathesoni.
Male. — Length, about 4.5 mm.; wing about 6 mm.
Rostrum, palpi and antenna; dark brown. Head dark.
Pronotal scutum and the collate dark brown; pronotal scutellum pale. Mesonotal
praescutum with three broad, reddish-brown confluent stripes, the humeral regions
cephalad of the lateral stripes pale; scutellum pale. Pleura pale, indistinctly striped
with bro%vn. Halteres pale, the knobs dark brown. Legs with the coxje and trochanters
pale; femora light brown; remainder of the legs broken. Wings subhyaline, un-
spotted; stigma lacking; veins brown. Venation: almost as in G. mathesoni with the
following details different: R2 very oblique and apparently contiguous with the tip of
Rl ; R2-\-J not angulated before the middle of its length and without a faint spur of r
at this point; petiole of cell M2 much longer, one-half longer than the fused portion
of Cul and M.
Abdomen light brown. Male hypopygium generally similar to that of G. mathe-
soni, differing as follows: The bifid pleural appendage is very similar in the two
species, in the present species with the needle-like tip of the longest arm abruptly pale.
The long, sinuous appendage in mathesoni is here represented by two, the longer of
which is pale throughout, flattened, the long tip acicular and almost straight; the
shorter appendage is flattened, before the tip a little expanded, with a long, slender,
curved black-tipped apex. Near the base of these pleural appendages is a flattened
subtriangular lobe which is covered with an abundance of short setae; in G. mathesoni,
this appendage is very small, cylindrical, with but few setae and with a distinct finger-
like spinous lobe on one side. Penis-guard distinctly trifid at its apex, the lateral black
spines directed almost caudad, setigerous at their bases; a shorter median pale lobe.
Habitat. — Colorado.
Holotype, <J , Longview, June 24, 1916 (E. C. Jackson).
Type in the collection of the United States Biological Survey.
90 Journal of Entomology and Zoology
tJenus Phyllolahis Osten Sacken.
Phyllolabis lalifolia, sp. n.
General coloration light gray; wings pale gray, the stigma pale grayish brown;
R2+3 shorter than RS alone; cell 1st M2 short; male hypopygium yellow with the
foliaceous appendage of the eighth sternite very broad and but indistinctly bifid at
its tip.
Male. — Length about 6.5 mm.; wing, 7.5 mm.
Rostrum dark brown, heavily gray pruinose above; mouth-parts reddish brown;
palpi dark brown. Antenna; moderately elongate, dark brown throughout, the flagellar
segments long-oval, provided with venticils that are but little shorter than the seg-
ments. Head light gray with an indistinct black median line.
Pronotum rather large, heavily light gray pruinose. Mesonotal prxscutum
brownish gray pruinose without distinct stripes; pseudosutural foveae black, short-
triangular; tuberculate pits not evident; remainder of the mesonotum gray pruinose,
the scutellum more brownish. Pleura clear light gray. Halteres pale. Legs with
the coxae and trochanters pale brownish yellow; remainder of the legs dark brown,
the bases of the femora paler. Wings pale gray; stigma rather indistinct, pale
grayish brown; veins dark brown; Sc and the abortive vein behind Cu more yellowish.
Venation similar to P. clai'iger but R2+3 shorter, less than R3 alone; veins R2 and R3
more divergent, R2 at the wing-margin being distinctly closer to R1 than to R3 ; cell
1st M2 shorter, especially the outer deflection of MS.
Abdomen brown, sparsely gray pruinose. Hypopygium light yellow, including
the pleurites and pleural appendages. Genitalia similar to P. clavigrr, differing as
follows: outer angle of the pleurite much longer, projecting conspicuously beyond the
pleural appendages; dorsal pleural appendage not slender and strongly bent at mid-
length but very broad and flattened, roughly subtriangular with the base narrowest.
Foliaceous appendage of the eighth sternite very broad and flattened, widest at the
base, thence wlili the sides almost parallel slightly expanded at the distal end, the
caudal margin of this leaf-like lobe slightly concave, feebly or indistinctly notched
medially.
Habitat. — Oregon.
Holotype, <J , Forest CJrove, March 2%, I9I9 ( F. R. Cole).
Genus Tricypliona Zetterstedt.
Tricyt>hona sfiarsi/>unila. sp. n.
Close to T. srptentrionalis Bergr. ; median prcsculal stripe split by a pale line;
wings subhyaline, the costal region more yellowish; r-m connecting R-f-^5 and Ml +2.
Femalf. — Length, 7.5 — 8.8 mm.; wing 9.2 — 11 mm.
Rostrum very short, transverse, dark brown, sparsely gray pruinose, the anterior
margin with a row of a few long yellowish bristles; mouth-parts and palpi dark
brown. Aniennx dark brownish black, the basal four or five segments enlarged and
very crowded as in this group of species. Head dark brown above, the front and a
narrow margin around the eyes and across the anterior part of the vertex light gray.
Mesonotum very high and gibbous. Mesonotal prsrscutum light grayish yellow,
with three dark brownish stripes, the median stripe split by an indistinct pale capil-
Pomona College, Claremont, California 91
lary line that is more distinct in front; the sides of the median stripe are nearly par-
allel; lateral stripes narrow, their anterior ends subacute; scutum with the lobes
marked with brown; scutellum light gray. Pleura dark brown, gray pruinose.
Halteres pale yellowish brown, the knobs dark brown. Legs with the coxae brown on
the outer face; trochanters dull yellow; femora and tibise dull yellow, tipped with
dark brown; tarsi dark brown, the base of the metatarsi paler. Wings subhyaline,
the costal and subcostal cells more yellowish; stigma oval, dark brown, paler
distally; sparse brown clouds along the cord, at the fork of R-f-\-5, along the outer
end of cell 1st M2 and, less distinctly, at the base of the sector; veins dark brown, Sc
more yellowish. Venation: The distance between Sc2 and the origin of the sector
shorter than the straight portion of the sector alone; Rs angulated and spurred at its
origin; upward deflection of /?/ slightly oblique, inserted in Rl rather far before its
tip, so that Rl-\-R2 is greater than the deflection of R2 alone; petiole of cell R4 short,
about one-fourth longer than r-m; r-m inserted between R4-\-5 and Ml-\-2; petiole
of cell Ml longer than this cell.
Abdomen dark brown; valves of the ovipositor reddish brown, strongly com-
pressed, slightly upcurved at the tip.
Habitat. — Oregon.
Holotype, 9, Hillsboro, April 1, 1919 (F. R. Cole).
Paratype, 2, Corvallis, May 14, 1917 (Moulton).
The type is much larger than the paratype but undoubtedly refers to the same
species. The fly is closest to T. septentrionnlis Bergr. (Alaska) in its spotted wings
but may be distinguished by the colorational and venational details as described above.
Subfamily Tipulinse.
Genus Tif<ula Linna;us.
Tipula mallochi, sp. n.
Belongs to the submaculata group; close to 7". submaculata Lw. ; male hypopygium
with the horns of the tergite short, outer pleural appendage not bifid, gonapophyses
short, eighth sternite with two powerful decussate bristles.
Male. — Length, 15 mm.; wing, 17 — 17.4 mm.
Female. — Length, 20 mm.; wing, 18.5 — 19 mm.
Frontal prolongation of the head brown, more yellowish above; palpi pale brown.
Antennae bicolorous, the flagellum with the basal enlargement of each segment black,
the remainder light yellow, on the apical segments a little more infuscated. Head
yellowish brown with a sparse grayish bloom; a capillary dark brown median line.
Mesonotal prasscutum dull brownish yellow with four rather narrow reddish
brown stripes, the remainder of the dorsum yellowish. Pleura pale yellow, whitish
pollinose. Halteres pale, the knobs dark brown. Legs with the coxae pale whitish
yellow; trochanters yellow; remainder of the legs darker. Wings pale gray, the
base of the wings and the costal region more yellowish; stigma brown; a brown
cloud at the origin of the sector; tip of the wing indistinctly darkened; obliterative
area before the cord in the base of cell R2.
Abdominal tergites dull brownish yellow, on the sixth to ninth tergites dark
brown; the caudal margins narrowly, the lateral margins more broadly, silvery;
92 Journal of Entomology and Zoolog)'
segments two to five with a narrow longitudinal brown sublateral streak; sternites
brown, the caudal margins of the segments pale. Hypopygium generally similar to T.
submaculata, differing as follows: Ninth tergile with the lateral horns very short and
broad, the tips acute, not long and tapering as in suhmaculala; outer pleural appen-
dage short and broadly flattened, the apex subtruncated, with a few coarse setigerous
teeth, in submaculata this appendage is more slender, tapering to the acute point, at
about midlength on the outer margin with a prominent spine to produce a bifid appear-
ance; gonapophyses broad and flattened at the base, the slender tips short, not long
and sinuous as in submaculata ; eighth sternite with a pair of strong reddish fused
bristles that are decussate, in addition to the smaller seta;. In the female, the sixth
and seventh tergites are dark brown, the ovipositor acute, the tergal valves being
especially long and slender.
Habitat. — Illinois.
Holotype, $, Alto Pass, Union County, June 5, 1919 (Alexander).
Allolopotype, 9.
Paratopotypes, 4^9; paratypes, 20^ 9, Makanda, Jackson County, June 4, 5,
1919 (Alexander and Malloch) ; 5 (J 9 , Dubois, Washington County, June 3, 1919
(Malloch).
Type in the collection of the Illinois State Natural History Survey.
Tipula malloclii is common in the "Ozark" region of southern Illinois during
early June, when it flies with other species of the genus as T. submaculata Lw., T.
tuscarora Alex., T. translucida Doane, T. morrisoni Alex., 7". mingtve Alex., T.
umbrosa Lw., T. ftavoumbrosa Alex., 7". fuliginosa Say, and, in proximity of low wet
cliffs, with T. ignobilis Lw.
Notes on Pacific Coast Pycnogonids
\V. A. HILTON
The specimens reported on at this time were obtained at Laguna Beach in the
summer of 1920. Their collection was more or less incidental to other littoral explora-
tions. There Is also included a list of forms obtained at other times and at other
places, chiefly during the same summer at Pacific Grove.
Pallene calijorniensis, Hall.
Two of these were collected at Laguna Beach.
Lecythorhynchus marginatus. Cole.
Twelve specimens collected at Laguna Beach from among mussels, under rocks,
among algae, etc. One specimen was dredged of San Diego in 1916. Thirty-four were
collected on the land side of Catalina Island at the Isthmus in quite a different type of
locality from that which is usual. At this place there were few red Algae but
masses of a rather fine brown rock-weed. On these plants, hydroids and bryozoans
were quite abundant. Many more might have been collected if there had been time.
Among Algae in front of the Hopkins Laboratory at Pacific Grove IS specimens
of this species were found. One was collected at the "Big Tide Pool."
Ammothella ttiherculata. Cole.
Twenty specimens found in front of the Hopkins Laboratory at Pacific Grove.
One found at low tide in the "Big Tide Pool." None found at Laguna this season.
A. bi-unguiculata, Dohrn, far. calijornica, Hall.
Twelve of these obtained at Laguna Beach under stones. Three specimens at the
Isthmus, Catalina Island.
A. spinosissima. Hall.
Seven specimens collected at Laguna Beach. Two obtained at Pacific Grove in
front of the laboratory.
Tanystylum intermedium. Cole.
Twenty-five specimens from Laguna Beach.
Clotenia occidentalis, Cole.
Ten specimens from Laguna Beach. Sixtey-three specimens from in front of the
Hopkins marine station, some were found on plume hydroids and among Algae.
Halosoma viridintestinalts. Cole.
We usually find a number of this species at Laguna Beach but none were found
this season. At Pacific Grove 68 were collected from masses of fine bryozoans from
floating timbers.
Amoplodactylus erectus. Cole.
Specimens of this species may be obtained at Balboa among tubularian hydroids,
a hundred or more were collected from this locality this year and one from Anaheim
Landing with palm hydroids.
A. califortiicus, Hall.
Three specimens from Laguna Beach, 6 specimens from the Isthmus, Catalina
Island, 1 specimen in front of the laboratory. Pacific Grove.
Pycnogonum stearnsi, Ives.
Seven specimens from Laguna Beach, 1 specimen from Pacific Grove.
(Contribution from the Zoological Laboratory of Pomona College)
Uca. musica. Railib.
Drawn by J. Caldwell from specimens obtained by Caldwell and Miss W. Diirani
at Balboa mud flats durini; llie summer of 1920. Tbis is the first record of a tiddler
crab in our region. The male is shown with the large claw. Sometimes the large
claw was on the right, sometimes on the left. Specimens brought to the laboratory in
moist sand made their burrows and lived all summer. In spite of the larger claws of
the males they gave way to the females when in each other's way.
Lepidopia myops, Stimp.
From Laguna Beach. Drawn by Joseph Caldwell
Kremila anatoga, Stimp.
Common sand crab of Laguna Beach.
Drawn bv Howard Lorbeer.
The
Nervous System
and
Sense Organs
BEGINNING WITH THIS ISSUE A SERIES OF ARTICLES
WILL RUN FROM NUMBER TO NUMBER
WITH CONTINUOUS PAGING
Bv WILLIAM A. HILTON
I. Plants
One of the common properties of living things is irritability.
All living substance reacts, responds to stimuli, whether they come
from the outside or from within. Transmission of stimuli is also
a common property of living matter.
Plants are sensitive to many sorts of stimuli without much
indication of organs of special sense. Only in certain cases are
there tissues for the transmission of the effects of stimulation and
central organs for coordination and control seem to be entirely
lacking.
In unspecialized organisms, both plants and animals, the sur-
faces are sensitive to many sorts of stimuli without special organs
for their perception. The whole surface or the whole body may in
a general way be sensitive. If there are special parts associated
with special stimuli, there are no histological features to indicate
them. This diffuse perceptive capacity is more characteristic of
plants than animals, yet some animals are of this type, and many
plants have structures which are truly organs of sense, and in some
cases special tissues for the transmission of the effects of stimu-
lation.
In certain parts of most plants there are areas of surface
where the perception of stimuli takes precedence over the pro-
tective or other functions; such surfaces may be called sensory.
Certain cells or cell groups in plants which have perception as their
chief or only function may be called sense-organs, even though
they may not be responsible for sensation in the psychological
sense. So far as we know, plants have developed sense organs
only in relation to a few forms of external stimulation, such as
those of contact, shock or jar, gravity or static and photic or light
stimuli. So far as we can tell, the real act of perception, so-called,
always takes place within the living substance, mainly or entirely
in the solid portions, or in the ectoplast.
Tactile pits occur in the outer walls of some surface cells. The
cell walls are thin at these points, which are just over the sensitive
protoplasm within the cells. These pits are usually confined to the
sides of tendrils which may come into contact with surfaces.
Darwin fir.st determined that tendrils can be stimulated only by
contact with, or friction against, solid objects, not by the impact
of water.
Tactile papillae, knobs and hairs occur on various parts of
plants, such as staminal filaments. Parts of flowers which exhibit
movements are often stimulated by means of hairs or knobs.
Movements of parts of insectivorous plants are initiated bj^ means
of special sensory structures, such as hair, knobs, or spines.
Plants respond to light in general without special organs of
sense, but it is probable that the epidermal cells of many leaves are
4 NERVOUS SYSTEMS AND SENSE ORGANS
arranged in such a way as to favor the reception of light waves.
This is, of course, not alone for sensation, yet sensation may be
an important function. Some epidermal cells bulge considerably,
especially in the velvet-like leaves of tropical forests. Such eleva-
tions make it possible for the cells to perceive photic stimuli, even
when their surfaces are wet. Sometimes a whole cell bulges in a
lens-like manner; sometimes the wall is thickened like a little lens,
and by these methods the rays of light are brought to a focus upon
the inner sensitive protoplasm. In many plants the whole unper
epidermis is developed as a light-perceiving or photic epithelium.
Also at times the margin or some definite locality has cells espe-
cially adapted to focus and receive rays of light. Such cells alone
or in groups are conical with rounded tips, the apex of each has its
wall thickened or almost biconvex. Such so-called ocelli have been
proved to condense the light more effectually than the ordinary
surface cell.
Stigmata or eye spots are found in certain plant spores and
among the flagellates, .such as VoIvhx. Enf/leiia. etc. In Kiqilena
the light-perceiving ability is confined to sensitive protoplasm near
the pigment spot. The eye spot or pigment therefore acts as a
light-screen.
Geotropic movements of plants are remarkable. The plants of
high organization especially seem sensitive to the stimuli of gravity.
Certain cells of roots, stems and leaves are provided with movable
starch grains. It has been suggested that the movements of these
starch gi-ains bring about changes for growth and movements ap-
propriate to the needs of the plants.
Transmissions of stimuli take place within cells from the points
stimulated to more distant portions, but they cannot well be deter-
mined. When the sensory and the reaction organs are more widely
separated the conduction is more obvious. In plants there are but
few examples of transmission at a distance, for in many cases of
marked movements in plants the sen.sory areas immediately adjoin
the motor tissue. In other cases the transmission is at a greater
distance. The velocity of transmission in plants is much lower than
in animals. Heliotropic and geotropic stimuli are .said to require
five minutes to travel two millimeters, traumic stimuli : 1-2 cm. per
minute to 1-2 cm. a second. In case of the .sensitive plant the trans-
mi.ssion is 30-100 mm. per second.
Besides the transmission of impulses through the protoplasm
of the coll there is the necessity for transmission from cell to cell.
No special pathways have been clearly determined for the first in
plants, but protoplasmic threads traverse the whole thickness of the
cell walls. It is questionable whether there are special structures
within plant cells for the conduction of .stimuli. Strands between
cells have l)een interpreted by .some as the jiathways of the effects
of stimulation. There is no central organ of coordination known
PLANTS 5
and no distinction is needed between afferent and efferent pathways.
The only instance known of special tissues for the conduction
of impulses is in the sensitive plant group and here it is quite defi-
nitely proved that living tissues are not necessary for the conduc-
tion of impulses and are in no sense comparable to the conductive
tissues of complex animals.
NERVOl'S SYSTEM AND SENSE ORGANS
II. Protozoa
In Amoeba, there seems to be no portion of the surface more
sensitive than others. The exoplasm is a general sensory organ.
Experiments by Hyman '17, with toxic substance show that a
local region of increased susceptibility exists along the axes of each
pseudopodium from it^ distal to its proximal end, the distal end
being more suscejitible. The youngest and most vigorous forms
are most susceptible.
According to several investigators, the exoplasm of Amoeba is
like a tough skin and this in part at least acts as a sensory area.
The more fluid endoplasm may become quite rigid under stimulation.
The changes in Amoeba which are the causes of amoeboid
movement and behavior originate within the Amoeba and external
stimuli do not act directly to produce those physical alterations
which result in movement, but they act through the jn-otoplasm of
the Amoeba. The reactions of Amoeba are similar to the reflexes
of more complex forms involving reception of stimuli, and the con-
duction of internal changes leading to response, but sensation, con-
duction and movement are not differentiated.
If one side of an Amoeba touches .some object it may move
away from the .source of stimulus. Jennings has found that when
touched the animal does not usually move directly awav from the
side stimulated, but merely in some other direction. If the anterior
edge is touched this part stops and contracts while the current
turns to one side at this point, so that the animal moves at an angle
with its former direction. If the advancing edge of an Amoeba is
touched it withdraws and a new pseudopodium is sent out else-
where. Sometimes Amoebae react positively to .solid bodies, they
may also under various sorts of .stimuli thrust out many pseu-
dopodia at once or draw all into a compact mass. Amoeba reacts
not only to mechanical but also to chemical, temperature, light and
electrical stimuli. The direction of movement in negative reac-
tions is not determined entirely by the position of the stimulating
agent. Other .stimuli may have already altered the character of the
protopla.sm, for exanii)le the moving Amoeba is temporarily dif-
ferentiated, having two ends different and the sides differing from
the ends. The.se and i)erhaps other internal factors have a large
part in the determination of movement.
It is impo.ssible to explain how Amoeba alters its own metabolic
process. If Amoeba is capable of self stimulation then this might
suggest that living sub.stance has a psychic quality which is pos-
sessed by all protoplasm. If this is not accepted for sim|)ler organ-
isms it would be hard to accept it for the cells of the cerebral cortex
of man and all would be referred to present or past conditions of
external or internal environment.
There is no clear evidence that Amoeba has memorv. The
PROTOZOA 7
nearest approach to a suggestion of it comes from the observations
of Jennings upon an Amoeba which attempted to devour one
smaller. The ingested specimen escaped its captor, the larger
reversed its movements and followed the smaller and again took
it in. The behavior of the larger might seem to be partly deter-
mined by its earlier experience, but this might also be explained by
a purely physical stimulus of a direct character.
Any elements of psychical qualities which Amoeba might pos-
Fig:. 1. Neuromotor systems and sensory systems of Protozoa. A, B. Dif-
flug'm showing the effects of stimulating the ends of the pseudopodia.
Verworn. C. Neuromotor system of Euplotes, Yocum. The moto-
rium is dark, strands to the orgmelles and to the cirri shown by
lines. D. EugJcua showing eye spot near gullet and flagellum. Kent.
E. Gonium showing eye spot above. Mast. F, G and H. Neuromo-
tor system in Diftlodbiinm after Sharp. The dark lines show the loca-
tion of the chief parts of the system. F. Ideal section of the whole.
G and H. Views from side and mouth end. I. Stem of Varicella after
Delage et Hevouard, the contractile portion shown in d-irk, the conduc-
tive part in lighter.
8 NERVOUS SYSTEM AND SENSE ORGANS
sess are not capable of demonstration or proof. All that we can see
is that if there are any elements of consciousness they must be of a
very vague and elementary nature.
All forms of protoiilasm have the property of irritability and
there is usually also involved a certain degree of conductivity, but
these are not always possible to measure or clearly determine. \'er-
worn has made a study of conductivity in the elongated thread-like
pseudopodia of some rhizopods. In studying the changes which
take place in the long pi'otoplasmic extension of DitHiujcu the re-
sults of stimulation may be directly observed. A weak stimulation
at the end of the pseudopodium causes a slight wrinkling of the
smooth surface, a stronger stimulus causes more swellings and more
distant ones on the slender appendages. Fig. 1, A, B. The extent
and rapidity of the wrinkling of the .surface is in direct response to
the strength of the .stimulus applied. Other species of rhizopods
gave similar results. The decrement of the intensity and rapidity
becomes greater with the distance from the point of stimulation
until the wave of e.xcitation is obliterated. This is of course in
sharp contrast to the conduction of a nerve fiber which normally
conducts excitations without perceptible decrement of the intensity.
An organ for the control of amoeboid movement has been sug-
gested, a centro.some or blejiharoplast from which strands radiate
to all the i^arts of the body which are concerned with locomotion,
l)ut no recent proof of this suggestion has come to my attention.
According to Hyman the nucleus in Amoeba plays an important
part in amoeboid movement, as is shown when the nucleus is re-
moved.
Ciliate Protozoa such as Paramoecium, Steiitor, VorticeUa,
etc., have much more complicated reactions than Amoeba becau.se
of their more complex structures, but the stimuli to which they
resjiond are not much more complex or varied. The cilia are often
highly specialized and localized; some coordination must be neces-
sary. Cilia in general have been described in various ways as asso-
ciated with small granules at their ba.ses and strands from these
granules have been described as penetrating into the cells, in some
cases at least to be associated with a body of nuclear or cytoplasmic
origin.
In 1880 Englemann found fibers in Stijlotu/chia to which he
assigned a nervous function. Nere.sheimer, 1903, found similar
fibers in Stoitar. and a number of others have described such
structures without always being clear as to their function. Sharp,
1913, considers an elaborate .system in Dijihulhiium which he calls
a "neuro-motor apparatus.' ' From a well-marked central body or
"motorium" strands of sub.stance were found going to the cilia
and to various parts of the body in a complex manner. Fig. 1, F,
G, H. Yocum, 1918, describes and figures a neuromotor system in
Ei( plates, developed from the ectoplasm. Fig. 1, C. It consists of
PROTOZOA 9
strands running from the motorium to sensitive areas, to the
membranelles and to the long anal cirri. There are also strands
connected with frontal, ventral, and marginal cirri, although these
are not connected with the motorium. These cirri are irregular
in their movements while the anal cirri are used chiefly in loco-
motion. These last as mentioned, have definite connections with
the motorium. Yocum traces the homology of the motorium
with the blepharoplast of many forms. This is the coordinating
structure which serves to regulate anterior and posterior regions
Fig. 2. Neuromotor systems of Flagellates. Nuclei and neuromotor appa-
ratus mostly shown by dark lines or masses. A-D. Origin of ble-
pharoplast from the nucleus in Xaegleria. x 1040. E. Flagellate after
Robertson, x 1200. F. Trypanoplasma after Martin. G. Trichomitus
after Kofoid and Swezy. x 800. H. Cercomonas after Wenyon. I.
ChilotnaHti.r after Kofoid and Swezv. x 318.5. J. Trichoni/mpha after
K. and S. x 1-50. K. Crithida, after McCulloch. x 1440. L. Leidu-
opsis, Kofoid and Swezy. x 200. M. Giardki, Kofoid and Christiansen.
x 2.550.
10 NERVOUS SYSTEM AND SENSE ORGANS
of the body. The basal granules of cilia, cirri, and membranelles
are considered as secondary rather than primary structures. In
ciliates the connection between neuromotor apparatus and cilia is
not clearly established, but there is some indication that there may
be connection.
In many flagellate protozoans the flagellum has been described
as springing from a center or blepharoplast. A very primitive type
of neuromotor apparatus is described by Wilson, 1916. The flagel-
lum arises from a blepharoplast which grows out from the central
karyosome (Fig. 2, A-D). The blepharoplast is connected with
the karyosome by a rhizoplast.
In other forms the blepharoplast may be composed of one or
more granules which may or may not be connected with the nucleus.
The basal granule of the flagellum may have a double function of
being a basal granule of the flagellum and also a division center for
the cell. In some forms the two functions are separated in two
granules. In some a number of granules surround the blepharo-
plast or may be derived from it. These migrate backwards and
come to form the ])arabasal lx)dy which may in some cases be at-
tached by a number of fibrils to the blepharoplast. This parabasal
body is interjn'eted as an acces.sory kinetic reservoir. A further
elaboration of this structure is the chromatic rod of some species.
Various types of flagellates with their internal connections are
shown in Fig. 3. One of the most complex conditions we find in
Ginrdia, Kofoid and Christianson, 1915. This is a binucleate organ-
ism equivalent to two flagellates, each containing one nucleus and
one blepharopla.st at the end of a single axostyle, three flagella and
a half or whole axostyle, depending upon the stage of the organism.
Two blepharopiasts are connected by cross commissures and are
anterior. The lateral flagella cross the middle line. The blepharo-
piasts are joined to the nuclei by rhizoplasts and also to the para-
basal body lying along the axostyle. "Each organism has its own
neuromotor apparatus, but due to the crossing of the fibers between
the blei)haroplasts the two organisms are unified. (Fig. 2, M.)
According to Yocum and others the motorium of ciliates is
homologous with the blepharopla.st of flagellates. According to
Dobell the blepharoplast of the jirotozoan is homologous with the
end knob and the axial filament of the metazDan sperm, whose func-
tion is to provide for the locomotoror activities of the cell. The.se
structures are akso homologous with the centrosome of resting cells.
It seems probable that other strands and coordinating centers
may be found in protzoans in addition to those already described.
This tyi)e of .system for control or coordination is not in any sense
homologous with that of Metozoa and in no .sen.se does it lead to
flevel()|)meiit of the nervous .system of more comi)iex forms. From
what has already been .said it is probable that methods of coordina-
tion are not at all alike in Protozoa and Metazoa; in fact it may well
PROTOZOA 11
be that the method in rhizopods may be of quite a different charac-
ter than in the more specialized Infusoria and Mastigophora.
It is quite interesting that the neuromotor apparatus is derived
from the ectoplasm. This corresponds to the probable conductive
tissues in the protoplasm of plants and suggests a comparison be-
tween the origin of these parts, with the origin of the nervous sys-
tem of Metazoa from the ectoderm.
Special sense organs in Protozoa are rare. In certain forms
there are eye spots or masses of pigment as in E}iglena, Fig. 1, D.
and there are also eye spots or sensory areas in such forms as
Goniiim. Fig. 1, E.
LITERATURE
A few only of the more valuable or suggestive references are given.
Bovard, J. E.
1907. The Structure and Movements of Condvlostoma patens. Univer-
sity of California Pub. Zool. 14, pp. 343-368, pi. 34. 21 text fig.
Biitschli, O.
1887-9. Protozoa. Bronn Class u. Ordn. des Tierreichs.
Delage et Herouard
1886. Traite de Zoologie Concrete, t. i. La Cellule et Protozoaires.
Dobell, C. C.
1909. Researches on the Intestinal Protozoa of Frogs and Toads. Q.
Jour. Mic. Sc. v. 53, no. 210, pp. 201-277. 1 text fig. pi. 2-5.
1909. Chromidia and the Binuclear Hypotheses: a review and criticism.
Q. Jour. Mic. Sc, vol. 53, no. 210, pp. 279-326. 25 text figs.
Englemann, T. W.
1880. Zur Anatomie und Physiologic der Flimmerzellen. Pfluger's Arch,
f. d. ges. Phys., pp. 505-35. pi. 5.
Hyman, L.
1917. Metabolic Gradients in Amoeba and their relation to the mechan-
ism of Amoeboid Movement. Jour. Exp. Zool., vol. 24, no. 1, pp.
55-99. 14 figs.
Janicki, C.
1911. Zur Kenntnis des Parabasal apparatus bei parasitischen Flagel-
laten. Biol Centrbl., t. 31, pp. 321-30. 8 figs, in text.
Jennings, H. S.
1904. Contributions to the Study of the Lower Organisms. Carnegie
Inst. Wash. Pub. 16, pp. 7-256.
1915. Behavior of the Lower Organisms. Columbia Univ. Press,
pp. 1-366.
Kofoid, C. A., and Christiansen, E. B.
1915. On Binary and Multiple Fission in Giardia muris Grassi. Univ.
Calif. Pui). Zool., vol. 16, pp. 30-54, pi. 5-8. 1 fig. in text.
Kofoid, C. A., and Swezy, O.
1919. Studies on the Parasites of the Termites III. On Trichonympha
campanula Sp. Nov. U. Calif. Pub. Zool., vol. 20, no. 3, pp. 41-98,
pi. 5-12. 4 text figs.
1919. Studies on the Parasites of the Termites IV. On Leidyopsis
sphaerica. Gen. Nov. Sp. Nov. Univ. Calif. Pub. Zool., vol. 20,
no. 4, pp. 99-116, pi. 13-14. 1 text fig.
12 NERVOUS SYSTEM AND SENSE ORGANS
1920. On the Morphology and Mitosis of Chilomastix mesnili (WenyonJ.
A common Flagellate of the Human intestine. Univ. Calif. Fub.
Zool., vol. 20, no. 5, pp. 117-144, pi. 1.5-17. 2 figs, in text.
.Machado, A.
1913. Sobre Ociclo evolutive de Schizolystis spinigeri N. Sp. Mem. Osw.
Cruz. t. V. fac. I, pp. 5-15, Estampas 1, 2 and 3.
Martin, C. H.
1910. Observations on Trypanosoma congeri. Part I. Divisions of the
active form. Q. Jour., Mic. Sc, vol. 55, no. 219, n.s. pp. 485-496.
pi. 21. 1 text fig.
Martin, C. H. and Robertson, M.
1911. Further Observations on the Caecal Parasites of Fowls with some
reference to the Rectal Fauna of other Vertebrates. Q. Jour. Mic.
Sc. vol. 57, no. 225, n.s., pp. 53-81, pi. 10-14.
Mast, S. O.
1916. The Process of Orientation in the Colonial Organism, Gonium
pectorale and a study of the Structure and Function of the Eye
Spot. Jour. Exp. Zool. vol. 20, no. 1, pp. 1-17.
Minchin, E. A.
1909. The Structure of Trypanosoma Lewisi in relation to Microscopical
Technique. Q. Jour. Mic. Sc. vol. 53, pp. 755-808.
1911. Observations on the Trypanosomes of the Little Owl (Athene
noetua), with remarks on the other Protozoan Blood-parasites.
Q. Jour. Mic. Sc. vol. 57, no. 226 n.s., pp. 141-185.
MeCulloch, I.
1915. An Outline of the Morphology and Life History of Crithidia lep-
tocoridis. Sp. Nov. Univ. Calif. Pub. Zool. vol. 16, no. 1, pp.
1-22, pi. 1-4. 1 text fig.
1919. A Comparison of the Life Cycle of Crithidia with that of Try-
panosoma in the Invertebrate Host. Univ. Calif. Pub. Zool. vol.
19, no. 4, pp. 135.
Porter, A.
1910. The Structure and Life-History of Crithidia melophagia (Flu),
an Endoparasite of the Sheep-ked, Melophagus ovinus. Q. Jour.
Sc. vol. 55. no. 218 n.s.
Sharp, R. G.
1914. Diplodinium ecaudatum with an Account of its Neuromotor ap-
paratus. Univ. Calif. Pub. Zool. vol. 13, pp. 43-122. pis. 3-7.
4 figs, in text.
Swezy, O.
1916. The Kinetonucleus of F'lagellates and the Binuclear theory of
Hartmann. Univ. Calif. Pub. Zool. vol. 16, no. 15, pp. 185-240.
58 figs, in text.
Verworn, M.
1913. Irritability. Yale Univ. Press, pp. 1-246.
Visentini, A.
1912. On the Morphology of the Leishmania of Italian Kala-Azar. Q.
Jour. Mic. Sc. vol. 58, n.s., pp. 353-370. pi. 19-20.
Wenyon, C. M.
1910. Some Observations on a Flagellate of the genus Cercomonas. Q.
Jour. Mic. Sc. vol. 55 n.s., no. 218, pp. 241-250. 19 text figs .
Wilson, C. W.
1916. On the Life-history of a Soil Amoeba. Univ. Calif. Pub. Zool.
vol. 16, no. 16, pp. 241 292. pi. 18-23.
Woodcock. H.M.
1906. The Haemofiagellates. Q. Jour. Mic. Sc. vol. 51 n.s.. no. 197, pp.
233-331. 64 text figs.
Yocom, H. B.
1918. The Neuromotor Apparatus of Eunlotes Patella. ITniv. Calif.
P.,l> 7„."1 vol. 18, no. 14, pp. 337 39fi. pi. 14-16.
SPONGES
13
III. The Sponges
The only activities of sponges which are in any way suggestive
of sense organs or a nervous system are those connected with the
water currents which enter and leave.
The currents are caused by collar cells distributed in the vari-
ous chambers. These flagellate cells cause the continuous move-
ments of the liquids under ordinary conditions. The flagella of
these cells are connected with basal granules or blepharoplasts in
each case and in some, connections are also made with the nucleus.
Fig. 3, I, J.
Lendenfeld, 1885-7, has described sensory cells and ganglion
cells in sponges, Fig. 3, E, F, G, but Minchin, 1900, and others
believe there are no true nervous elements. No modern work has
suggested the possibility of nerve cells or sense cells in Porifera.
Parker, in 1910, describes elongated spindle-shaped cells ar-
ranged like irregular sphincters around the gastral cavity, oscu-
lum, etc. Structurally they have the appearance of a primitive
TTTnxnni
FiR. 3.
Structures from sponges. A. Dermal membrane of a sponge seen
from the exterior. Membrane pierced by six pores, three of which are
partly closed by pore membranes. After Wilson, after Parker. B, C,
D. Three stages in the closure of the membrane pore. After Wilson,
after Parker. E, F, G. Sense cells and nerve cells, (?). After von
Lendenfeld. H. Two stages in the development of a muscle cell as the
first stage in the development of the nervous system. Diagram after
Parker. I, J. Collar cells from sponges. After Robertson, x 1,000.
K. Transverse section of the base of an oscular collar of a sponge
showing the cavity surrounded by a sphincter of myocytes, spicules
outside. Modified from Parker.
14 NERVOUS SYSTEM AND SENSE ORGANS
kind of smooth muscle fiber. As a result of their contraction the
opening into the sponge is lessened or closed.
Wilson, 1910, describes membranes covering the subdermal
cavity and containing pores. This so-called membrane is composed
of an e.xternal portion and is believed to be syncytial. There are
two somewhat independent devices for the closure of pores, the pore
membrane and the pore canal sphincter. The closure of the pore
canals is dependent upon the sphincter-like band of cells on the
wall of the canal. These cells are in every way comparable to a
primitive form of smooth muscle-fiber. They are in contact with
the water passing into the canal and seem capable of direct stimu-
lation. The pore membrane is less muscle like and is perhaps of a
more primitive type.
Parker, 1910 and 1919, considers the sponges as an important
group in illustrating the most primitive condition of the nervous
system of metazoans. Muscle cells the independent effectors, as
illustrated by the sphincters of sponges, were the first neuromuscu-
lar organs to appear. The special receptors in the way of sense-
cells were next to appear in certain coelenterates while in other
forms more complex, the adjuster or central organ was added.
LITER.ATURE
Bidder, G.
1896. The Collar-ceils of Heterocoela. Q. Jour. Mic. Sc. n.s. vol. 38,
pp. 9-43. pi. 2.
Lendenfeld, R. Von
1885. Das Nervensystem der Spongien. Zool. Ang. Bd. 8, pp. 47-50. 2 fig.
1887. Synocils, Sinnesorgane der Sponpien. Zool. Anz. bd. 10, pp. 142-
145. 2 text fig.
Minchin, E. A.
1900. Sponges. A Treatise on Zoology edited bv E. R. Lanlcester. Part
2, eiiap. 3.
Parker, G. H.
1909. The Origin of the Nervous System and its Appropriation of Ef-
fectors. Pop. Sc. Mo. vol. 75, pp. 56-64.
1910. The Reactions of the Sponges with a Consideration of the Origin
of the Nervous System. Jour. Exp. Zool. vol. 8, pp. 1-41.
1919. The Element-irv Nervous System. Monog. Exp. Biol. pp. 1-229.
53 illus.
Robertson, M.
1911. The Division of the Collar Cells of the Calcarea Heterocoela.
Q. .Jour. Mic. Sc. n. s. Vol. 57. no. 226, pp. 129-139, pi. 19.
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JOURNAL OF
ENTOMOLOGY AND
ZOOLOGY
VOLUME XIU. 1921
PUBLISHED QUARTERLY BY THE
DEPARTMENT OF ZOOLOGY OF POMONA COLLEGE
CLAREMONT, CALIFORNLA. U. S. A.
CONTENTS OF VOLUME XIII
Volume XIII, Number 1
Gemmell, Ella
A Note on a Local Mcnilur ol llu-
Family Psychodidac, 1.
Campbell, Arthur S.
Littoral Opliiurans at Lacuna
Beach. 2.
Moles, M., and Moore, W.
A List of California Arachnida.
Fscudoscorpionida. 6.
Hilton, W. A.
TIk- .Nervous System and Sense
Organs. Part IV. 15.
Index to Volume XII. 27.
Volume XIII, Number 2
A List of California Arachnida.
IL I'edipalpida. M. Moles. 11.
III. Scorpionida. F. A. Cox. 12
IX. Solpugida. J. Ncsbet. 14.
Campbell, Arthur S.
.\oles (111 the Sense Organs of
Some .Asteroids, 16.
Hilton, W. A.
.N'crvous System and Sense
Organs. V. .U.
Volume XIII, Number 3
A List of California Arachnida
\'. I'lialangida. L. Myers. 19.
\'l. .Xcarina. F. Cox. F. Jahraus.
W. Moore. 23.
Hilton, W. A.
.Wrvous System and Sense
Organs. '\"I. 40.
Volume XIII, Number 4
A List of California Arachnida.
\'ll. Araneida. M. Moles. I. John-
ston. .?').
Campbell, Arthur S.
Ophiurioidca of the West Coast of
North .'\mcrica. -16.
Hilton, W. A.
Nervous System and Sense
Organs, Vll. 55.
INDEX TO VOLUME XII
Arachnida, D, 11. 1'^, y>.
Araneida. 39.
.■\steroids. 16.
Campbell, A. S.. 2. \h. -td.
Coclcntcrata. 15.
Cox, F. A., 12. 15. 2,i.
Eyes. 16.
Flat-worms. .54.
Fly, 1.
Gemmell. E.. 1.
Hilton. W. A.. 15. ,i4. 45. 55.
Jahraus. P.. 2.!.
Johnston. I.. i9.
Mites. 2i.
Mules. M.. 6. 11, 39.
.Moore. W.. 6. 23.
Myers, L., 19.
Xemertinea. 49.
Nervous System. 27. 34. 49. 55.
Nesbet. .1.. 14.
Ophiurians, 2. 46.
Fedipalpida, 11.
Pscudoscorpions. 6.
Psycliodidae. 1.
Ronnd worms, 55.
.Scorpionida. 12.
Sense organs, 16.
Serpent stars. 2.
Solpugida, 14.
Spiders. 39.
Ticks, 36.
Wliip-seorpions, 11.
h NOV 17 1939 ^^■
VOLUME THIRTEEN NUMBER ONE
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
MARCH, 1921
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT o/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
A Note on a Local Member of the Family Psychodidae —
Ella Gemmell 1
Littoral Ophiurans at Laguna Beach — Arthur S. Campbell - 2
A List of California Arachnida Pseudoscorpionida —
M. Moles, W. Moore 6
The Nervous System and Sense Organs, Part IV —
W. A. Hilton 15
Index to Volume Twelve -----27
Entered Claremont Cal..Post.Office Oct. 1, 1810, as second-tlass matter, under Act of Cor.gress ol
March S. 1878
Journal of Entomology and Zoology
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riareinont, California, U. S. A.
A Note on a Local Member of the
Family Psychodidae Dip
Ella Gemmell
A number of specimens were collected about a drain in a house in Claremont.
As they had not been seen here before and as there was no known standing water
near, it was a question as to where they had passed their earlier stages. All the
specimens were collected in or near the outlet to the small sink. Afterwards it was
found that the cesspool was nearly filled and a new one had to be made, after this
the flies disappeared until once again when the cesspool was filled the flies appeared
in the house.
These specimens were determined to be PsychoJa cinerea Bks. this being the
synonjTn for P. pacifica as described by Kincaid. The specimens agree in most all
parts to Kincaid's description. The specimens were 2 mm. in length. The figure is
from a male.
Littoral Ophiurans at Laguna Beach
ARTHUR S. CAMPBEI.I.
During the summer of 1920 specimens of all species of (jpliiurans previously
known to exist near Laguna were obtained.
Several stations were found to be constant with the various species, some for
adults, and others for voung. Several are limited to very special habitats. Two
are limited to but one locality each, while O. splailata is found abundantly in almost
all stations examined. For the first time O. maculosus was found inshore under
stones; previously this species had been known only from kelp holdfasts.
The excellent plates for this paper are the work of Miss E. Keyes, a student
in Pomona College.
Ophiodermatidae
No dental papilla. Buccal papillae numerous. Two or four genital bursa: in
each interradius.
Ophioderma paniimeitsis Liitkin.
Add. Hist. Oph., 2, p. 193. 1S59.
Large. Arms, three or oflener four times diameter of disc. Mouth papillar
and teeth small. Arm spines numerous, flattened, lying close to arm. Color dark
brown above, lighter below, the arms encircled by pale bands.
Young in Macroeyslh holdfasts. Adults in rocky tidepools among Finns and
green algz, ranging up to middle littoral tide pools. Common.
Opiiiotryptus mttiulosus Clark.
Third Laguna Report of Pomona College, p. 64. 1915.
Small. Disc covered with swollen plates concealed by rough granules. Upper
arm surfaces more or less covered by granules. Oral shields except madreporite,
adoral and oral plates covered completely by granules continuous with above. Five
almost conical, subequal arm spines. Two tentacle scales. Color white, grey or
with disc marked with reddish granules. Pise in young is red, becoming marked
later only by a few red granules, and Anally dirty while in adult. Seventeen arm
joints. Arms one and a half times diameter of disc.
In Macrocystis holdfasts, washed inshore under loose rocks. Young and adults
intermingled. Rare.
OPHIOr.EPIDAE
No dental papillx. Three or six buccal papillar. Always i\vo genital bursa,
nisc notched. No tooth papillx.
Opiiioplntus fsmarki Lyman.
Bull. M. C. Z. 3, pi. 10, p. 227, pi. 5.
Medium sire. Arms nearly three times diameter of disc. Three arm spines.
Disc with plates on both surfaces. Disc and arms flattened. Color light or dark
brown, some blue-grey.
Young in Mmrtiiyslis holdfasts, in calcareous sponges and among red alga; in
lide-zone. Adults in rocky tidepools among Funis and green algr; in sand un<ler
loose rocks. Abundant.
Pomona College, Claremont, California 3
Amphiuridae
One to five mouth papillse. Arms arising from ventral side. Two genital bursae.
Amphiodia barharae Lyman
III. Cat. M. C. Z. Harvard, 8, pt. 2, p. 17, pi. 3. 1875.
Medium size. Arms twelve or more times diameter of disc. Oral papillae six,
equal and regularly arranged. Teeth. No tooth papillae. Two short, flat tentacle
scales. Three tapering arm spines. Color yellowish or tinged with green.
Deep in sand at Balboa. Young in sandy pool on shells. Rare.
Ophionereis annulata Le Conte.
Proc. Acad. N. Sc. Phila., 5, p. 317. 1851.
Medium size. Arms about six times diameter of disc. Mouth papillas numerous.
Teeth. No tooth papillas. Three flattened, stout arm spines. Color light, arms dis-
tinctly banded.
Young in sponge masses. Young and adults among beds of Mytitus, Lepas and
Mitella; in sand under loose rocks, and in rocky tidepools among Funis and green
algae. Common.
Ophiocomidae
Mouth papillae. Teeth. Arms arise from ventral side of disc. Two genital
bursae. Mouth shields small or medium.
Ophiopteiis papulosa Lyman.
111. Cat. M. C. Z. Harvard, 8, pt. 2, p. II. 1875.
Large and coarse. Arms three or four times diameter of disc. Disc completely
covered above by stout stumps. Few mouth papilla?. Five flat, blunt arm spines.
Color deep brown, arms often faintly banded.
In rock ledges with ground shell or sandy bases. Associated with the echinoid
S. purpuratus Stimp. around the mouth region of which there is often a member of
this species of opliiuran. Rare and restricted.
Ophiothricidae
Plates on upper side of arms small. No oral papillae. Tooth papillae. Few
buccal papillae.
Upliiothrix spiiulala Le Conte.
Proc. Acad. N. Sc. Phila., 5, p. 318. 1851.
Variable size. Arms five or six times diameter of disc. Disc covered with
thorny spines. No mouth papillae. One tentacle scale. Seven long arm spines.
Color greenish brown, red or yellowish. Arms with orange bands. Mouth usually
whitish. Some have red discs. Color variation in this species is extraordinary;
apparently there is no uniformity.
In Macrocystis holdfasts; in rocky tide-pools with Fucus; in mussel-beds with
Mytilus, Lepas and Mitella. Young also found in calcareous sponge masses, and
4 Journal of Entomology and Zoology
among red algi in rock tidepools. Very common ; the most abundant species found
at Laguna Beach.
ASTEROPfnTIDAE
Teeth and mouth and teeth papilla; spiniform, indistinguishable. Arms re-
peatedly divided.
Gorgoncephalus eucnemis M. & T.
Syst. der Aster. Braunschwig. 1842.
A specimen of this species was obtained several years ago, on a line about 160
faths. some distance from the Laguna shoreline. It measures 130 mm. across the disc.
(Contribution from the Laguna Marine Laboratory of Pomona College.)
Fig. 1. Ophioderma panamensis Lulkiii.
Fig. 2. Ophiocryptus maciilosus Clark.
Fig. 3. Ophioplocus esmarki Lyman.
Fig. 4. Amphiodia barbarae Lyman.
Fig. 5. Ophionereis annulata Le Contc.
Fig. 6. Op/iiopteris papulosa Lyman.
Fig. 7. Op/iiol/irix splculata Le Conte.
(All figures are X2, and of the dorsal surface.)
A List of California Arachnida
This list is compiled from already published hut scattered papers. Many of these
are local records of specimens and new species collected hy many students through a
number of years and determined for us for the most part by Banks and Chamberlin.
As numerous earlier papers in this Journal have taken up the distribution of local
forms only a hint of this will be given. There are included in this list records other
than local. If the distribution is general some indication is given. A few hints as to
characteristic features are given when possible. The family characteristics are com-
piled by the aid of the works of Banks, Ewing, Comstock and several others. In
order to save space the literature references are given in abbreviated form at the end
of each section, especially as there are a number of papers and lists already published
which give this material in great detail.
I. PSEUDOSCORPIONIDA
M. Moles and W. Moore
Cheliferidae. Evidences of segmentation of thorax in some species. Serrula
attached all its length to finger of chelicera. Spinneret long and slender. Flagellum
absent. Tarsi of legs one-jointed. Tarsal claws short and thick, split on some of
the feet.
Chelifer cancroiJes Linn, about buildings, oak, sycamore trees, Claeremont, mts.
C. fuscipa Bks. Calif.
C. scabrisculus Simon. N. Calif, to Claremont.
Chtlanops ohlongus Say. Palm springs, Brown's flats.
C. vaWdus Bks. From Lake Tahoe.
C. paUipet Bks., under stones Claremont, I-os Angeles.
C. dorsaVtt Bks., Lake Tahoe and San Francisco.
C. acumitiatui Sim. Maraposa, Claremont, Laguna Beach.
C. lagunar Moles, Two eye spots. Claremont.
C. paludis Moles, Claremont.
C. serratus Moles, No eye spots. Clavate hairs saw-like edge.
Aiemnus hirsutus Blcs. Lagiina Beach near ocean.
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(!aryt>us lalijornuus Bks, t'lulcr stones I.aguiia Beach. Also Palo Alio and San
Nicolas Island.
PsfuJngrypus hicnrnii Kks. Sliasla Springs.
Ideobsiid.xr. Spinnerel long. Serriila attached only at base. Carapace not
divided.
tdfohisium magnum Bks. Ml. Shasta. Four eyes.
/. Ihrevenrii Simon, Four eyes. San Francisco to Claremont.
Urrnnfus ohicunii Bks. Lake Tahoe and Claremont.
Pomona College, Claremont, California 9
Obisiidae. Spinneret small knob. Serrula attached only at base. Carapace not
divided.
Ohisium macilentum Simon, Claremont-Mt. Shasta.
Bhtlirus calif ornicus Bks., S. Calif.
B. magnus Ewing. Shasta Springs.
Linn Syst. Nat. ed. 12, 1767. Ann. Ent. Soc. Fr. 1878. Jour. N. Y. Ent. Soc. 1895.
Jour. Ent. Zool. June 1914. Jour. Ent. Zool. Dec. 1911, V. 3, p. 633, 1914, 6, p. 818,
p. 6, Nn. 4, p. 87, V. 9, 1917, p. 26, V. Canad. Ent. 1893, p. 67, also 1891, p. 165.
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IV. Coelenterata
HydrozoA Polyps. The structure of fresh water Hydra has
been studied with reference to the nervous system for some time.
One of the earlier papers was by Korotneff, '76, who recognized
nerve cells. Later work was by T. J. Parker, '80, Rouget, '81, and
Schneider, '90. This last author determined a network of ganglion
cells to be present. Zoja, '90-'92, finds structures in Hydra which
he believes are nervous elements because they take special stains
and according to him have connections with the epithelial muscle
cells and with nematocysts. These cells are similar to the ganglion
cells described and figured by Schneider.
Citron, '02, in Syncoryne a compound hydroid, finds spindle-
shaped sense cells especially in the end knobs of the tentacles. Gang-
lion cells with three or four processes are found in various parts of
the body while bipolar ganglion cells are found in the coenosarc.
WoM, '03, determined that hydroid polyps have a nervous sys-
tem, partly of sense cells, partly of ganglion cells. The processes of
the latter are more or less joined. The sense cells are primitive
intra-epithelial. There is quite a complex network of fibers and
cells on the body and tentacles, quite a concentration also on the
manubrium. Long strands of the plexus run the whole length of
the polyp. There is a less abundant entodermal plexus.
Hadzi in '09, used the isolation method with Hydra, also sec-
tioning methods. He found a plexus of nerve cells all over the
surface of the body and tentacles ; these were chiefly triangular
shaped cells. He distinguishes bipolar and tripolar cells as well as
some multipolar forms ; the first are sense cells. He shows anasto-
moses at various places. The greater part of the system is an ecto-
dermic network. He says that it is not appropriate to speak of
neurones, for the cells are directly connected by protoplasmic proc-
esses, and Hydra is too far from the type in reference to which the
neurone concept was established.
The palm hydroid Conimorplia, which is more favorable than
Hydra tor experimentation, has Iseen studied by Torrey '04, Parker,
'17, and others.
The reaction systems of coelenterates are cilia, nettle cells,
mucous glands and muscles. In this genus, mucous glands and cilia
are not important. Nettle cells are apparently independent of ner-
vous control, a condition not true of Hydra if we accept the work
of several investigators.
There are six sets of muscles in Corymorplia; two of these are
entodermic, the circular muscle of the stalk and the circular muscle
of the proboscis. When anesthetics which control nervous tissues
are used, these two muscles remain capable of acting". This proba-
blv shows that these muscles are not under control of the nervous
16
NERVOUS SYSTEMS AND SENSE ORGANS
system. The four other muscles, the longitudinal muscles of the
stalk, proboscis and the two sets of the tentacles, are quicker in
their action and are controlled by anesthetics. These are probably
supjilied by sense cells and the nerve-net.
Stimuli applied to any part of the normal animal may be trans-
mitted to distant parts; strong stimuli are transmitted to more dis-
tant parts than weak ones. The nervous transmission is probably
limited to the ectoderm. Although the nervous system is very
primitive, reactions much like a true reflex occur, as Parker has
pointed out. When a proximal tentacle is strongly stimulated ad-
jacent tentacles resjwnd and the proboscis may turn to the stimu-
lated point.
Fin:. 4. A. General plan of the nevrous .system in Hydra. B. Nervous system
of Artiiiia. Diapramatic. C, D. Hydroid jellyfish showing position
of eye spots. Mayer. E, F. Otocysts of hydroid jellyfish, Mayer.
G. Otocyst and eye spot, Mayer. H. Hydroid medusa with eye spoU:
at base of tentacles. I. Tentaculocyst, Mayer. .1. Eye spot with
biconvex lens, Mayer. K. Tentacle and eye spot, Mayer. L. Diagram
of the nervous system of a hydroid jellyfish. Wall of the bell cut
away on one side showinp section of manubrium and gonad. M. Ten-
taculocyst and eye spot, Mayer. N. General plan of the nervous sys-
tem of a scyphozoan. Diapramatic.
Many parts of the jiolyp are (juitc independent of the re.st of
the body, as may be seen when the hydranth has been removed; the
stalk will shorten and even localize a stimulus apjilied to one side.
The hydranth is not necessary for coordinated resjionse. Neither
COELENTERATA
17
is the stalk necessary for reflex movements of the tentacles and the
proboscis.
The neuro-muscular organization of Corymorplia is diffuse and
in no sense centralized.
Hydroid Medusae. Although the nervous system of medusae
is of the difi'use type, there are concentrations of the network at
certain places. In Gonionemiis there is a double ring of nerve cells
and fibers about the margin of the bell. Hyde, '02, mentions a third
ill-defined ring; this might be considered to be a part of the diffuse
network or plexus which is found over the surface of the sub-
umbrella. In addition to the two main marginal bands, there are
concentrations of nerve cells and fibers following the four radii of
the bell, and the manubrium has some concentration of nerve cells
and fibers.
Although the nerve ring is usually double, sometimes it is not
divided. The nerve tissue is between the ectoderm and the muscu-
lar tissue. In some forms the peripheral system is but poorly de-
veloped with only a few nei've cells scattered beneath the surface.
Fig. 5. Nerve cells from various coelenterates from a number of sources.
A. Nerve cells from Hydra, Schneider. B. Nerve cells and sense cells
from jelljiish from Kasseanow. C, E. Nerve plexus from Siphono-
phora, Schneider. D. Sense cells and nerve cell in Hydra. F, G. Nerve
cells and fibers in the epithelium of Hydra, Wolff. H, I. Nerve cells
from actinian, Hertwig. J. Nerve cells from Cerianthus, Grosley.
K, L. Nerve cells from actinian, Havert.
18 NERVOUS SYSTEMS AND SENSE ORGANS
In Lizzia. the Hertwig brothers, '78, found the tentacles
jrrouped, and at the base of these the nerve cord is swollen, due to
a concentration of ganglion cells. The suggestion has been made
that the two nerve rings have different functions; the upper one
connected with the sense organs, the lower being near the muscles
gives nerves to them.
Loeb found that if the bell without the nerve ring be placed
in five-eighths per cent NaCl or five-eighths per cent NaBr. it goes
on beating rhythmically, but small quantities of CaCl, or KCl or lx)th
added caused the bell to stop contracting. The bell would beat in
sea water if not for Ca or K, and possibly some other ions.
When two specimens of medusae are grafted together after the
nerve rings are removed, the two portions contract as one and not
from two centers of contraction.
Krasinska, '14, in Connarhia finds large and small ganglion
cells and two kinds of sensory cells. The ganglion cells are mostly
multipolar and in a sub-epithelial region nerve elements are also
found in the tentacles; large ganc^lion cells are found in the sub-
umbrella and small in the tentacles. The velum is enervated by
fibers from the inner nerve ring. She does not decide whether there
is a true nerve network because she found but few cases of anasto-
mosis. The large ganglion cells of the sub-enithelial plexus are con-
sidered to be motor, also the smaller ganglion cells of the tentacles.
In a hydroid medusa, Tiaiopi^is, Romanes found that the manu-
brium reaches over to a spot .stimulated by touch. Romanes found
that this movement continued after the margin with the nerve ring
was removed.
Loeb explains the coordinating movements of medusae In-
simple irritability and conductivity without attributing other spe-
cial functions to the ganglion cells exce|it those which occur in all
conducting protoplasm.
Yerkes, '02, determined that the medusa Guiiiiiucmii.^ has a
delicate chemical sen.se. All portions of the lx)dy except the velum
and exumbrella are sensitive to chemical and mechanical stimuli.
The tentacles are especially sensitive to nhotic stimuli. The inten-
sity of the .stimulus determines the (uiickness. duration and extent
of a reaction. Stimuli which affect s\ inmetrical ))oints of the body
uneoually have a directive influence noon the movements. Yerkes
concludes that the reactions of si'ecial Parts of Giinioiicmiis are not
dependent for their execution upon the functional activity of the
central nervous sy.stem. Irritabilitv is a property of all parts of
the animal except the iellv of the bell and the exumbrella surface,
bi't it differs widely in different regions.
As lyoeb suggests spontaneitv is not dependent UPon the central
nervous system but upon a hii^h degree of irritabilitv of cei'tain
parts of the margin of the bell, Thos" <"e<itiiens with the marginal
COELENTERATA
19
ring removed do not show spontaneous movements because insensi-
tive to other than strong stimuli.
Coordination is not dependent upon the function of any nerve
center, but upon the rapid transmission of an impulse.
Krasinska finds fibrillae within the ganglion cells of hydroid
medusae by means of the iron-hematoxylin method.
In Poiijorchis, Little 1914, there are two nerve rings, the lower
being the larger. All the cells are bipolar. Connections between
nerve cells and eye were not determined.
Work by Romanes '98 shows nervous connection between the
tentacles and also the manubrium.
The sense organs of medusae, marginal octocysts and eye spots
Fig. 6. A. Nerve cells from nerve rings of Gonionemus, Hyde. B. Eye spot
above lithocyst below tentacle base, medusa. C. Eye spot at base of a
tentacle of a hydroid jellyfish, Little. D. Tentaculocysts hydroid
medusa, Mayer. E. Tentaculocyst hydroid medusa. Haeckel. F. Ten
taculocysts from Trachimedusae, Mayer. G. Tentaculocyst hydroid
medusa, Haeckel. H. Section through sense organs, eye spots and
otolith of scyphozoan jellyfish. I. and J. Front and side views of
scyphozoan jellyfish sense organ. K. Simple eye of medusa Schewai-
koff. L, Section through more complex eyes of Anrelia. M. Marginal
notch and tentacle of AnrcIia from above. Eimer. N. Section through
marginal tentaculocyst of scyphozoan showing sense areas, dark.
O. Section through tentaculocyst scyphozoan, Hesse.
E., from Dahlgren and Kepner's Histology. G., M. and N., from Parker and
Haswell Zoologv, permission of Macmillan Co.
20 NER\'OUS SYSTEMS AND SENSE ORCIANS
are often found, but the two kinds are not usually in the same ani-
mal.
In Lizzia, the eye spots are found on the under side of the ten-
tacle, but in this form the tentacle is held up and its lower side
turned toward the litrht.
In Poliiorchis, Little '14, the eye spot is on the outer surface
at the base of each tentacle. In other naked-eye medusae, similar
conditions are found ; the eyes may be arranged about the margin
as in this form, or in groups to correspond with the groups of ten-
tacles.
In genus Tri()j)sis. there are eight marginal sense organs con-
sisting of an entodermal ocellus and an open fold of velum which
contains concretions. In Phopal'Dicnia the lithocy.>;ts are indo.sed.
In the Narcomedusae there are maririnal sensory clubs con-
taining concretions of entodermal origin. Romanes. '98, found that
if the bell of a hydroid medu.'^a was removed the contractions of the
bell cease, but the margin which contains the nerve ring continues
to contract as before the injury. Any iniurv of the umbrella
causes no change in the rhythm so long as the rerve ring is intact.
The conclusion from this was that the nerve rinar is a coordinating
center and one needful for rhvthmical conti'actions.
In many medusae, otocvsts or senorv clubs probably function
as static organs. In Anthroniedusae there are no otocysts. but
many have ectodermal ocelli on the bnses of the tentacles. Romanes
found that these had certain v'su^l functions. Medusae with them
were strongly attracted to light between the red and violet spec-
trum.
In some forms like nouf/aivrHlia. the tentacles are grouped
and to correspond to each tentacle at its base is an ocellus or pig-
ment spot.
In the Lentoinedusae there may be marginal sensory clubs and
there may be lithocysts of ec-t"dermic oriirin. In some forms such
as Landicca there mav be marginal sense clubs with no concretions
within and ectodermal ocelb at the b-^ses of the tentacles.
In OrrJiistonia pileiis Larson, there are four hundred dark
brown entodermal ocelli on the circular canal : each is provided with
an pctodermal lens.
ScYPHOZOANS. The marginal sense organs of this grouo arc
so marked as to be early recocrnized. Ehrenberg. 1837. was the first
to .sneak of these as organs of sense. The usual type is somewhat
as follows. At eight marginal notches we find two small tentacles
either side of a shorter hollow tentacle. This tentacle or tentaculo-
cvst contains otolyths and the ovfixn seems to be one of equilibrium.
I'non the surface of this tentnculncvst there mav he a special pig-
ment .spot or ocellus of rather simnle structure. In the little flap
above and also behind or below the short sensory tentacle there may
be special areas of cells which may have some sort of olfactory or
COELENTERATA 21
chemical sense. Both Eimer and Romanes published physiological
papers in 1877-1878 on work done several years previously which
seemed to show that jellyfishes had the power of conducting im-
pulses in a complex manner along their subumbrellar surfaces.
Taschenberg, 1877, was unable to find nervous structures and
considered that the muscles responded directly to stimuli without
the aid of a nervous system. The Hertwig brothers, 1878, clearly
demonstrated the existence of a nervous system in medusae.
Schafer, '79, found a network of nerve fibers in the subumbrella
lying between the muscular layer and the ectoderm, but did not de-
termine anastomosis. Somewhat later Schlater, 1891, believed he
had found the true nervous system in the marginal sense organs,
but a clear recognition of nerve cells was again made by Kassianow
ten years later. He found a nerve plexus in Lucernaria and Cra-
teroloplivs. In the latter, tripolar ganglion cells are also found.
He shows sense cells and ganglion cells in direct association with
epithelial cells.
Hesse, '95, in Rliizostoma shows the structure of marginal sense
organs in some detail and gives some indication of the nervous sys-
tem. Fibei's run from the eight marginal sense areas to a more or
less circular band which is somewhat poorly defined, and other
strands spread out over the subumbrellar muscular bands of the
jellyfish. The relation between cells was not clearly shown.
Bethe, '09, was able to prove that the nerve plexus in Rliizo-
stoma is a true network.
Romanes and others have shown that the bell of a jellyfish
could be cut in a most complex manner without preventing the pas-
,sage of a stimulus for a contraction wave.
If a single marginal body is stimulated, contraction waves
start both to the right and to the left of the stimulation until they
mingle and disappear.
If the center of the jellyfish is cut out and the margin deeply
notched, the tortuous pathway of tissues thus formed is capable of
carrying a contraction wave. If a jellyfish with one marginal sense
organ is cut in a spiral strip, a wave of contraction may be started
at the margin which will run the whole length of the strip.
A jellyfish cut so as to make two concentric rings with only
two slight connections between will carry the impulse from the outer
to the inner portion by this narrow bridge. If the jellyfish is cut
so as to form a long circular stretch, a wave may course for a long
period round and round the bell. Such a "trapped" wave has been
known to go for eleven days with no great decline in rate; or at the
rate at which it was traveling, it would have covered a distance of
four hundred and fifty-seven miles in eleven days, Parker, 1919.
The removal of the marginal bodies of a medusa causes the
movements to cease for a time, but it may be made to contract by
electrical or chemical .stimulus. Experiments by Bethe seem to show
22 NERVOUS SYSTEMS AND SENSE ORGANS
that although the muscle of the jellyfish is capable of direct stimu-
lation, it is not so sensitive as the nerve-net. Parker summarizes
the susceptibility to stimulation as follows: 1. Marginal bodies
most sensitive. 2. Nerve-net. 3. Muscles directly stimulated least
sensitive.
Mayer, 1917, concludes from his experiments with Cassiopea,
that nerve conduction is due to a chemical reaction involving the
cations of sodium, calcium and potassium. The proljable high tem-
perature coefficient of ionization of this proteid may account in some
measure for the high tention coefficient of the rate of nerve condi-
tion, which he finds is two and five-tenths as great as that of the
electrical conductivity of the seawater surrounding the nerve. His
observations do not support the "local action" theory. The rate of
nerve conduction is practically identical whether sea water is diluted
with 0.415 molecular mercuric chloride or with distilled water.
Corry, 1917, working with the same species found that regener-
ation takes place more rapidly on the half of the jellyfish in which
the sense organs were not removed. When sense organs are re-
moved and one half stimulated by electricity and the other insulated
half not stimulated, regeneration is more rapid on the activated
part. The experiments show that the rate of regeneration is but
one e.xpression of the general metabolic activity of the animal and
as such is subject to the influence of the nerve centers as are many
other functional activities. It is concluded as a result of experi-
ments that some chemical interchange between sense organs and
the surrounding tissue is necessary in order that the activity of
these structures shall be maintained at the highest state of efficiency.
Some sort of trophic influence is exerted in general metabolic
activities by the sense-organs. The structure of the nervous system
of this foiTTi makes it impossible to prove the existence of tropic-
nerve fibers as distinct from those of sensory or motor functions.
In Pehif/ia, Krasinska finds large and small ganglion cells in
as.sociation with sense cells. The large ganglion cells are considered
to have a motor and the smaller ones a .sensory function. There
are three methods of connecting the nerve plexus with the epithelial
surface. (1) Through peripheral processes of the ganglion cells.
(2) Through .sense cells. (.3) Through free nerve endings. No
direct proof of the enervation of the muscle fibers was established.
The tentacles have large and small ganglion cells, the cells are
deep in the muscular folds but in the outer eiiithelium is a fine
nerve-fibrillar area. Similar fiber masses are found in other parts
of the body and the nervous system ; these may correspond to a
"neuropile." Fibrillae were found especially in the branches of
the ganglion cells.
ACTINIANS. The reactions of the actinians have attracted at-
COELENTERATA 23
tention from qute early times; Milne-Edwards in his natural history
of corals in 1857 wrote :
"They enjoy a highly developed sensibility, not only do they con-
tract forcibly on the slightest touch, but they are also not insensible
to the influence of light. But no nervous system or organs of
sense are to be discovered in them." In these early times there
were, however, some vague suggestions of ganglia and nerve chords
in Actinia, but no confidence was placed in them. Huxley, in his
elements of comparative anatomy of 1864 says : "The nervous sys-
tem has at present not been determined in them." Alexander
Agassiz, in his seaside studies of 1871 says: "Only a few pigment
cells found at the tentacles are sense organs."
Schneider and Ritteken, 1871, state that the chromatophores
are organs of sense, compound eyes.
J. D. Dana in his Corals and Coral Islands, states that "they
sometimes possess rudimentary eyes."
Duncan, 1874, describes in some detail the structure of the
"eyes" of actinians. He also recognizes a plexus or network of
nerve fibers and cells under the epidermis, and remarks that the
difi'use nature of the nervous system is what might have been
anticipated.
The first rather complete account of the nervous system was
by the Hertwig brothers in 1879-80. They recognized sensory cells
in the epithelial layers and under the epithelium and next to the
muscular layers of both ectoderm and entoderm a layer of nerve
fibers and cells. The sen.sory cells when .stimulated carry impulses
to the nerve cell layer and this in turn to the muscles beneath them.
Nerve impulses from the ectoderm to the entodermal muscles were
supposed by them to pass over the exterior to the oesophagus and
from its inner end to the entodermal musculature. They considered
the body of the sea-anemone to be rather uniformily supplied with
nervous tissue except at the oral disc where in the ectoderm the
cells were concentrated in a sort of center. WolflF, 1904, and Gros-
ley, 1909, in the main accepted Hertwigs' suggestions but they
placed the concentration of the nerve fibers in the wall of the
oesophagus and not in the oral disc.
Kassianow, 1908, in Alcyonaria, believed the disc to be the
center of an individual member of the colony and Liedermeyer,
1914, although his observations were of sections alone, was of a
similar opinion from his study of one of the Pennatulacea.
Havert, 1901, on a sea-anemone by means of the Golgi method,
maintained a diff"use nervous system for actinians. This author also
believed that the ganglion cells, so-called by the Hertwigs, were
really motor cells which receive impulses from sensory cells and
then transmit them to muscles, a condition more like that of the
central nervous svstem of forms with a refiex arc. This author also
24 NERVOUS SYSTEMS AND SENSE ORGANS
showed a direct connection between ectoderm and entoderm, a con-
clusion which Parker, 1917, and Parker and Titus, 1916, have
shown on both anatomical and physiological evidence.
Von Heider, 1877, was of the opinion that the mesenteries of
some actinians miyrht contain nervous elements. Wolff, 1904, and
Kassianow, 1908, were of the opposite opinion but a number of
investigators seem to have shown that Von Heider's opinion is the
right one, among them Hickson, 1895, Ashworth, 1899, Kiikenthal
and Proch, 1911, and Liedermeyer, 1914.
In recent years Parker has given this group considerable atten-
tion and some of his conclusions will be employed in the following
discussion. There is also a paper on the histology of actinians by
Sanchez, 1918, but in this the nervous system is not considered very
extensively.
The effector systems of sea-anemone are mucous glands, ciliated
epithelium and muscles. Although nematocysts are considered by
some to be under control of the nervous system, there is good evi-
dence that they are independent of it. The only system under the
control of the nervous system is the muscular. By means of exj^eri-
ments it was learned that the bases of the anemones were esijecially
sensitive, but nervous transmission may be accomplished from
almost any poi'tion of the ectoderm to its longitudinal mesenteric
muscles. By several experiments it was proved that the trans-
mission might be by means of almost any narrow bridge of tissue,
proving quite conclusively that the transmission is by a nerve-net.
Many muscles responded at some distance from the point stimu-
lated and in some cases muscles were capable of responding directly
to a stimulus; whether these mu.scles were also under the control of
the nervous system at other times was not clearly established in
every case. In the acontia, however, there seemed to be no inter-
mediation of nerve impulses in the response to stimuli. Connections
from ectoderm to entoderm was proved in many cases. In con-
necting the ectodermic and entodermic system the lips and oesopha-
gus seemed not as imjjortant organs as other parts of the body.
Although the system of the actinians is diffuse there is some
degree of specialization. If the tentacles are stimulated by a nu-
trient fluid the oesophagus gapes by contraction of the transverse
mesenteric muscles, while weak acid causes a retraction of the oral
disc by means of a contraction of the longitudinal me.'^enteric
muscles. The two kinds of re.spon.se suggest independent receptors
and relatively independent transmission tracts.
In the tentacles the ectodermal surface is more receptive than
the entodermal; if there is a nervous .structure in the latter it is
probably very simple. The tentacles are complete neuro-muscular
organs and may react quite independently of the polyp, as shown
when severed from the body.
COELENTERATA
25
Parker has measured the rate of transmission of the nerve
impulse in sea-anemones at 21- centigrade. It was found to be
from 121-146 mm. a second.
Kassianow, Parker and others have studied the nervous system
and reactions of colonial forms. There seems to be little evidence
of any nervous coordination in colonial polyps, each polyp in Renilla
for instance when stimulated by contact seems to react independ-
ently of the rest. Although the common flesh which supports them
may bring about like changes in several or all of the members of
the colony, the zooids are not centers from which impulses pass to
other parts.
The peduncle and rachis are probably permeated by a nerve-
net which extends from the zooids of the colony.
Ctenophora. The first observations on the nervous system
of this group were by Pschschiltz, 1829, and later by Mertens, 1833.
One of the first complete summaries of the general structure of the
Fig. 7. Two-thirds of an elonj^ate ctenophor, Mayer. B. Enlarged portion of
sense organ of elongate ctenophore. C. Diagram of a ctenophore,
Mayer. D. Sense organ of ctenophore from side showing connections
with the eight ciliary glands. E. Same as D from above. F. Nerve
plexus of a ctenophore, Hertwig. G. Apical sense organ of a cteno-
phore after Hertwig. H. Diagram of a ctenophoi-e, Hertwig. H. and I.
View of apical sense organ of a ctenophore showing its relation to the
ciliary bands. H from the side, I from above. J. Coenoplana from
above showing apical sense organ, Korotneff. K. Coenoplana sense
organ in section with associated ganglia, Abbott.
I., J., from Parker and Haswell's Zoology, permission of Macmillan Co.
26 NERVOUS SYSTEMS AND SENSE ORGANS
nervous system was by Hertwig, 1880. A subepithelial nerve plexus
with the bipolar and multipolar cells has been described and figured.
Bethe, '95, also describes and figures a network of nerve cells and
fibers in ct^nophores.
The characteristic aboral sense organ was first described by
Edwards, 1841. At a lat«r time Chun, 1878, describes and figures
it in detail showing the little otocyst with its group of calcium
crystals supported on four bands of fused cilia like a little table,
with each tip of the leg coming into relation with two of the eight
ciliary bands.
This peculiar balancing organ has been considered in a way to
represent a central nervous system because of its reaction to the
ciliary bands. These bands seem not to be under the control of the
nerve cells and fibers, but some are of this opinion. The nervous
system then would not relate to the cilia, but in some way there
is a coordination of movement in the eight ciliary bands. That this
is not as simple as might at first seem is shown by the fact that the
effective stroke is in the op])osite direction from the wave of ciliary
action, so that the simple explanation of the movement of one cilium
affecting the next, like a row of tenpins, does not hold.
Bauer, 1910, found by gently touching the mouth region of a
ctenophore, that it .stopped its cilia. If vigorously .stimulated
its plates vibrate more actively for a short time. If the abt)ral
sense organ be removed the same reactions apply as before. He
concludes from this that the reactions cannot be ascribed to the
sense body but must depend upon the action of the diffuse nervous
system which although chiefly concerned with the nnuscles of the
cetenophore seems also to have an influence on the rows of swim-
ming plates.
Gothlin in a recent paper, 1920, on the study of ciliary move-
ments finds that the primary inhibition of the ciliary movement
is probably due to cilio-inhibitory nerves. Receptors at the surface
of the Iwdy transfer their impulses to the nerve-net. These in turn
transmit them to the end apparatu.ses which inhibit the vibrations
of the swimming plates, probably blocking the neuroid conduction
between them. There is an intimate connection between primary
and secondary inhibitory mechanisms. Both probably use the same
receptors, but the primary mechanism functions on impulses of
weaker intensity.
Abbott, 1904, who has studied the intere.sting worm-like Coelo-
phuKi has found a rudimentary nervous system with four ganglia
symmetrically disposed about the otolithic capsule. Just outside
the otolithic capsule in the angles formed by the intersecting tenta-
cular and sagittal jilanes are four large nerve ganglia that .send off
fibei"s to form a sort of diffuse peripheral .sy.stem and supply fibers
that cover part of the capsule as an enveloping sheath. Each gang-
COELENTERATA 27
lion is opposite the point of insertion of the cilia which support the
otolith. The cells of the nerve tracts and ganglia are large, tri-
angular and stain deeply with methylene blue.
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Medusa Gonionemus murbachii. Part I. The Sensory reactions
of Gonionemus. Am. Jour. Anat. vol. 6, no. 6, pp. 434-449. Part
II. The Physiology of the Central Nervous System. Am. Jour.
Phys. vol. 7, no. 2, "pp. 181-198.
Zoja, R.
1890. Alcune ricerche morphologiche e fisiologichi suU' Hydra. Dissert.
Pavia. 99 pp. 6 Taf.
1892. Die vitale Methylenblaufarbung bei Hvdra. Zool. Anz. Bd. 15,
pp. 241 2.
1892. Intorno ad alcum particolarita di struttura dell' Hydra. Rend.
Inst. Lomb. Milano, vol. 25, fac. 9, 13 pp, pi. 3.
1893. Sur quelques particularites de structure de I'hydre (systeme ner-
veux). Arch. Ital. Biol. t. 18, pp. 3.50-362, 1 pi.
(^ NOV 17 1939 i^:
VOLUME THIRTERP^ NUMBER TWO
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
JUNE, 1921
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT o/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
A List of California Ar.'vchnida
II. Pedipalpida, M. Moles 11
III. SCORPIONIDA, F. A. Cox 12
IV. SOLPUGIDA, J. Nesb(t - - - - - - - 14
Notes on the Sense Organs in Some Asteroids —
Arthur S. Campbell 16
Nervous System and Sense Organs
V. IV. A. Hilton - . - 34'
Entered Claremont, Cal., Post-OfBce Oct 1. 1910, as second-class matter, uoder Act of Congress of
March S, 1879
Journal of Entomology and Zoology
EDITED BY TUMONA COLLEGE, DEPARTMENT OF ZOOLOOY
Subscription $1.00 to domestic, $1.25 to foreign countries.
This journal is especially offered in exchange for zoological
and entomological journals, proceedings, transactions, reports
of societies, museums, laboratories and expeditions.
The pages of the journal are especially open to western ento-
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and Californian forms and conditions are particularly desired,
but short morphological, systematic or economic studies from
any locality will be considered for publication.
Manuscripts submitted should be typewritten on one side of
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Manuscri]its should be sent by express or registered mail.
Address all communications to
The Jol'rnai. of Entomology and Zoology
William A. Hilton, Editor
Claremont, California, U. S. A.
A List of California Arachnida
II. PEUIPALPIDA OR
WHIP-SCORPIONS
M. Moles
ScHIZOKATlDAE. Eyes wanting, caudal appendage short, unsegmented or knob-like
segment at end.
Trilhyreus pentapeltis Cook. Found rather commonly about Claremont, Laguna
Beach and farther south.
Tar.antvi.idae. The tailless whip-scorpions. Eight eyes.
Acaniho/ilirynus cnrnnatus. May be nearly two inches long. Calif, possibly
some specimens in the Pomona College collection may have some from the southern
part of the state but no clear record.
Proc. Ent. Soc. V. 4, p. 2^1. Jour. Em. Zool. V. 9, 1917, p. 1.
A List of California Archnida
A l.lSr OK CAI.II OKMA AKArllMDA
111 im St'dKIMOMDA
Krcil A. Cox
Bl THIDAE. Iriaiigtilar Menuim. Due or two spurs on cacli siile al liase of la~i
pair uf le^s. Three to Hvr lateral eyes on each side. Maud of chelae rounded, lin-
gers long. I'sually a spine under the sting.
i'roflfdes mex'uaniis. No spine under sting. Tecih on linger of palpus in man>
ohiique rows. Texas and Calif.
tniit iihilus i.innc
Saiil.i K:irli:ir:i an. I t ;ii;i
Mauds
-Iciulcr Icif)!
Tilyus tniuimanus Kks, Bucca Vista.
('rntrurus laliforiiinu Wood. Lake Tiile and Lake Co., Calif.
C fxiliiaudiis Wood. Lower Calif, and near San Diego.
ScoRPlovinxR. OnK one spur at hnsr of last tarsal segment of last pair of leg^
Pomona College, Clareniont, California 13
Diploifiilnis iiliilri. Texas nnd Calif. Twelve tn ciKliteen teelli on comli.
C'HAUTIDAK. Only two lateral eyes on each side.
Hroteas alleni W^ood, letitjili 1 to 1 ' j inches.
Vejoviu.m;. One spur each side of the hasc of the last tarsal segment of last pair
{)f lejis. Three lateral e\es on eacli siile. Sternum iisiialK' broader than long. \o
s.iine inider sting.
L'roiliiiius miirdax Trorcll. Hark colored, large claws, ("otnmon in Central and
N'orthern Calif.
.Iiiiirni tonus /'/ini-iulin lylns W'ooil. Rather hairy, red-hrown. San Hiego, Mojave
Pesert, Claremnnt. Common species.
I'ejov'u punclii'dlpi W \. Red-browii, strongly ridged claw. Oeath \'alley,
San Diego Co.
/'. Iiirsutii iiiidii Bks. San Bernardino Co. Red-hrown, 15 pectines. Length 1 '4
inches.
IlaJitius liirsulus Wood. Deserts of S. Calif.
C. Jour. Vm. \. 2, I9III, p. IS5. Ann. Mag. Nat. Hist. XVII, 1S76, p. II. Jonr.
.\c. Nat. Sc. Fhila. 1S63, pp. 3S7, 369, 372.
A List of California Arachnida
IV. sui.pri;inA
J. Nesbet
SOI.PUGIDAE.
Eremohales jiirmnnria Kocli. large specimen trom Brawlev. No spine* under
lihia in either sex.
E. faliforniia Sim. From Laguna Beach and Calif. Movable tin(jer of male con-
stricted from below near apical third.
Pomona College, Claremont, California 15
E. jormidabiiis Sim. Small spines under side tibia of palpus of male. Calif.
E. putnami. No spines on tibia of palpus of male. Calif.
Hemerotrecha californica Bks. Upper finger of chelicera wtih no teeth or mativ
small teeth. Pacific Grove to Claremont.
Ammoirecha californica. Lower finger of chelicera fine teeth bevond large teeth
at base. Broad dark band on middle of metatarsus of palpus. Calif.
Class des Galeodes 1879, p. 143. Ent. News 1903, p. 79. Jour. Ent. Zool. IX, p.
22. Proc. Acad. Nat. 1883: 3, p. 349.
Notes on Sense Organs in Some Asteroids
\Knn R ^. L'XMI'IIKII
Tlie sense organs of many species of starfish have been well studied during the
past fifty years by a number of competent observers. Among the earlier important
studies are those of Haeckel, 1S60; Wilson, 1862, and Hamann, IggS. Later work,
especially the more minute observations are the subjects of study of Cuenot, 1887,
and of Pfeffer, 1901.
Materials for this study include most of the common littoral asteroids occurring
at Laguna Beach. Representatives of six species, the members of three orders, were
examined. .Ml preparations were fixed in HgCI.. and double stained, first in hema-
to.Yylin and then in picro-fuchsin.
Eyes are placed at the terminus of each ray, and jusi proximal and ventral to
the terminal tentacle. In nearly all species they are well protected by a strong
circlet of heavy spines. They are mostly of a deep red color which is slowly
soluble in alcohol.
Viewed more closely the e>e-spot appears as a pad in which there are a number
of little depressions; these are the ocelli. Each presents a separate structure, the
whole eye-spot being merely a composite of many ocelli. The number of ocelli
varies greatly.
The histology of the ocelli in these forms has been disputed by several observers.
Most of the earlier workers believed that lenses are present. Cuenot, 1887, does not
accept this, but Pfeffer, 1901, indicates a lense in Asleropeitin miilleri. In some of
my preparations there is a little indication of an epithelial thickening bridging the
eye-cavity, but mostly the eyes show a clear and rather wide, open space freely in
communication with the exterior. These preparations indicate somewhat an inter-
mediate condition between the two figures reproduced from Pfeffer.
Cells forming the eye are of two types. The several reproduced from Cuenot'.-
paper, fig. 12, are pigment cells or sensory cells of the retina. They are surrounded
and supported by cells of a second type; the so-called supportive cells of Cuenoi
and others.
The comparative structure of sever.TJ eye preparations is figured. Tlie sup-
portive cells are well stained with fuchsin.
A sense organ in starfish was seen in Lintkia rolomliiir CJrey, among my prepara-
tions in ijie course of this investigation. It is probably a tactile organ. It is seen
in the ventral porlicn of the terminal tentacle, near the eye-spot. It consists of a
number of papillx extending over a restricted area of the tentacle. The papills are
pronounced and have a similar structure to (hose found in other forms. The>
follow through a small series of sections rather completely, showing constant form.
These may be like the so-called organs of taste described by Eimer, 1880.
(('.onlrihiiluin /ram llir Zonloi/iiiil l.ahnratory nj I'omnnii Collrgf)
BIBI.IOC.RAPIIV
CufNOT, I.. 1887.
Contribution a I'etude anaiomiipie des .'\sterides.
.Arch, de Zool. Exp. et Gen., 2f serie, vol. 5 his (supp.) p. 52-pl. 3, fig. 11-18.
Eimer, Tii. 1880.
Pomona College, Clarcmont, California
Neben Tastapparate bei tiii/rins mullitornis.
Arch. f. Mic. Anat. XVIII, pp. 34_^-346.
Haeckel, E
Ueber die Augen unci Ncr\'eii der See>tfrnc.
Zeit. f. VViss. Zool., 10, 1860. Tat. 11, pp. 183-190.
Ham.ann, O.
Beitrage zur Histologic der Echiiiodermcii, 2, Die Aster.
Anatomie u. Hist. Untersuclit. 7 pi. Jena.
Pfeffer, W.
Die Sehorgane der Seesteine.
Taf. 41, pp. 5-23-550. Zool. Jalnhucli. Anat. 14.
JOURDAI.V
Sur les Venx de I'Ast. rnliens.
Comptes-rendus de I'Acad. des. Sc. Tome 60, 1865, p. 103.
\\'ll.SON, N.
The Nervous System of Asteridse; with observations on the organs of sense.
Trans, of the Linnean Soc. 1862. vol. 21. pp. 107-123. 3 PI.
INDEX TO FUU'RES
Fig. 1. Ventral and lateral views of eve-pad I'isiistcr iiipilalns. showing general
relationship to terminal tentacle. .Xy.
Fig. 2. Ventral view of eye-pad of Orthiisin i/niiali'iui. X9.
Fig. 3. \'entral \'iew of e\'e pad of l^i.uistt'r .ot /nenfus. X9.
Fig. 4. \'entral view of eye-pad of .1 siniriii miiiiatii. .\9.
Fig. 5. N'entral view of eye-pad of l/imkui inlomhitii-. X9.
Fig. 6. Ventral view of eye-pad of .1 sliinpi'din rriniiifui. X9.
Fig. 7. Ocellus froin Orlliastn iionnlcnii to show general form. X350. Drawn by
camera lucida.
Fig. 8. Ocellus from Liiiikiii inlamliiar to show general features. X350. Camera
lucida.
Fig. 9. Ocellus from Asleiinn mitiiiitti. .\350. Camera lucida.
note the clear central margin of pit.
Fig. 10. Tactile organ from terminal tentacle of I.iinhiii lolombia
showing papillae and details. Camera lucida. .\350.
Fig. 11. Single sensory cell from l.iiiikui t olum/iiiir. Very greatly inagtiified.
Fig. 12. Sensory cells from Asterias nihrns showing plgmetit. Reproduced from
Cuenot. Osmic acid. Greatly magnified.
Fig. 13. General view of eye-pad of Jslci n^idiii criiii
lucida.
Fig. 14. Simple ocellus in an Aslnias. Supportive cells dark
Reproduced from Pfeffer. Diagramatic.
Fig. 15. A more complex ocellus from .1 stndpcclin mulleri.
features as above. From Pfeffer. Diagramatic.
CJeneral view,
Cieneral view
us. X350. Camera
Sensory cells lighter.
Note the lens, other
V. Flatworms
TURBELLARIA. Among the turbellarian flatworms those of the
Rhobdocoelida are the simplest. Bohming. 1890, describes and
figures a number of central nervous systems from Alioeocoeia such
as shown in Fig. 8. The ganglia are somewhat concentrated but
show right and left halves. Two or four pigment spots imbedded in
the brain substance may show but little indication of differentiation
into eyes.
Among the Acoela some have simple pigment spots for eyes
and some are without them. Statocysts are found in the center of
the ganglionic masses in some cases. Very often a well-marked
statocyst or otocyst may be seen in the center of the upper portion
of the animal, just between the pigment spots when they are pres-
ent. The brain is not very extensive in Acoela. It is usually recog-
nized as a small mass of cells surrounding the central statocyst.
Lohner in Pvlyclioenis gives about as complete account of the
nervous system as any. There is a central ganglion with a central
otocyst. Laterally there are two ganglia of nearly equal size. These
ganglia in cross section are nearly central in position while the
peripheral nervous system consists of longitudinal strands both
dorsal, ventral and lateral in position. Figure 8 shows the plan of
the nervous system as a whole.
De Quatrefages, 1884, and Peebles, 1915, and others give some
indications of the nervous system and .sense organs of these worms,
but not much in detail.
Many investigators have dealt with the Rhabdocoela. The
brain is a little more complex than that of the other groups men-
tioned but the whole s.vstem is compact and there are few longi-
tudinal cords from the brain region.
Some forms have from two to four simple eyes imbedded in the
brain. Sensory pits near the head end are found connected with
the brain in .some. Ott, '92, describes "dish-shaped" organs near
the dorsal surface of the body of Stenostoma. In this form the
ciliated pits are imbedded in the forward portions of the brain. In
other forms, thev seem to be entirely separate.
Schneider. '7.3, finds the lobes of the brain connected by a
double com'ni.ssure which surrounds the vascular system. Hallez,
"79, Ott, '92. and others find but a single commissure.
The fibrous jxjrtion of the brain or "punkt substance" is com-
posed of a fine network of fibers which some have thought was
made up of ana.stomosing processes, but the evidence is not clear.
Nansen, '87. does not believe in an anastomosis.
Some of the figures from the nervous systems of this group
show few branches. Probably more branches were present although
not recognized in ever.v case by the investigators.
FLAT WORMS
35
Fig. 8. Nervous System of rhabdocoelida. A and B. Brain with one and
two pairs of eyes of alloeocoelan flat worms, Bohming;. C. Nervous
system of an acoelan, Polychoerus, Lohner. D to H. Brains of Rhab-
flocoela. D. Oii'n^tniua. E. Prorhynchiis, after Vejdovsky. F. GaffiUa,
Bohming. G. Rhabdocoela nervous system, Bohming. H. Stennntoma,
Ott. I. An acoelan showing nervous system after Bohming.
36 NERNOUS SYSIKM AM) SKNSI. ()R(JANS
The brain consists of a rather broad Hat mass of nerve fibers
and cells occupying quite a large part of the forward portion of
the head end. Many nerves run out to the surface of the body and
two chief longitudinal strands run the length of the body. L'suallv
a number of commissures connects the two parts of the brain as well
as the two longitudinal strands. The number of these is .somewhat
variable in the different species and also in members of the same
species. In some forms at least, terminal fibers connect peripheral
branches at the margin of the body. Fig. 9. A, D. E. l'suallv two
eyes are found connected with the brain by short nerves, but in some
cases at least, such as in Sorocelis, as described in Seidl, 1911, there
are neurone eyes scattered over the anterior region of the forward
end.
Lateral extensions of the head end are often especially sensitive
and provided with abundant nerve cells. The eyes, simple or com-
ple.x have been well described and figured by Hesse, 1896. A sen-
.«ory cell or cells with expanded ends terminate in a i)igment cup
which aids in centering the light on the protoplasmic ends of the
sen.se cells. Fig. 9 F-H.
Very little has been done in analyzing the motor and sensory
components of the brain and nerves. Branches to the eyes and to
the surface of the body, especially the forward end of the body, are
undoubtedly sensory in nature. The brain has been divided by
some into an anterior and superior .sensory region and a posterior
and inferior motor portion. Some of the chief works on this grouj)
are by Chichkoff, '92; lijoma. '84; Lang. '81; Woodworth, '91;
Wheeler, '94; Voidovskv, '95; Hesse, '97; Micoletzkv, 1907; Weiss,
1910; Seidl, 1911.
Rina Monti, 1896, has studied the nerve terminations in the
skin of fresh-water planarians.
The Polycladida are usually considered as having a more com-
plex nervous system than the tricladids. but it is more concentrated.
As a rule there is a number of simijle eyes scattered over the for-
ward end of the lx)dy such as shown by De Quatrefages, 1844,
although in Pkniocera Lang, '82, shows rather concentrated eye
areas. In Leptoplana, the eye spots are scattered about in the
region of the nervous svstem, as shown bv Schmidt as earlv as
1862.
Although locomotion in planarian worms may in part be by the
surface cilia, the chief activities seem to be by means of muscles
of the body under the control of the nervous .system. Weak chemical
or tactile stimuli cause them to react positively. The resting worm
re.snonds less readily than the moving one. Some forms with much
more highly organized eyes react less well than others with simnler
eye spots. As a rule .strength of light is le.ss important in reactions
than the riimber of sen.sory elements in the eye, or the forme-
FLAT WORMS
37
habits and experiences of the animal. Headless forms respond to
light but less quickly. As a rule if the head and eyes of a planarian
are removed the headless portion reacts as before but much more
slowly. In marine flatworms where the ganglia are more concen-
trated in the head region and where there are fewer ganglion ceils
along the lateral cords, the activities of the headless worms are
much less perfect than in planarian worms of fresh water.
In the flatworms special cells of the ectoderm give rise to the
head ganglia. Later stages, or the development of the peipheral
svstem have been but little studied.
Fig'. 9. Nervous system of polyclad and triclad worms. A. Snycoelidium,
Wheeler. B, C. Head and tail ends of Sorocelis, Seidl. D. Brain and
head end of Pla)iaria bohmegi, Weiss. E. Planaria apitia, Micholetzky.
F and H. Eyes of Planarians, Hesse. I, J. Nerve endings in skin
Planarians after Monti. K. Brain and eyes of Lcptoplana, Schmidt.
L. Nervous system and eyes polycladid, Lang.
Kepner and Rich, 1918. have studied the reactions of the pro-
boscis of flatworms. In accordance with Monti, '97, and Steiner,
'98, they found that the ventral nerves are ganglionic and these
centers exercise control over the posterior parts of the body. The
middle branch from each of these ventral nerve trunks leaves the
ganglion that lies nearest the base of the proboscis and from here
enters it. When the proboscis is removed from the animal it
undergoes autoamputation. Without the control of the adjacent
.?8 m:r\()us system and sense organs
ganglia the proboscis in this way acts as a reflex organism. The
freed proboscis is able to carry out the three usual coordinated
muscular movements when the muscles are intact. The free pro-
boscis cannot determine food from other substances. The central
nervous system is necessary for this.
The eyes of turbellarians have been extensively studied by
Hesse, '96. In tricladids they consist of visual cells and pigment
or acsessory cells. These last inclo.se the enlarged ends of the
visual cells, the rhabdomes. The number of visual cells or retinulae
as well as the acces.sory or pigment cells differs greatly. Kepner
and Taliaferro, 'IG, found the retinulae to consist of three regions;
a lateral nucleus bearing region closely applied to the brain with
a nerve fiber extending into it, a middle region lens shaped, homo-
geneous and highly refractive, and the true rhabdome in the pig-
ment cup. Kejiner and Foshee, '17, compare the three regions
of the retinula with the rods and cones of vertebrates. The parts
show a close analogy if not homology with the myoid, ellipsoid and
rhabdome. The retinulae of both flatworms and vertebrates are
also of the inverted type. Taliaferro, 1920, has an important paper
on the reactions of Planaria to light. The species considered was
negative to light and turned itself accurately to horizontal rays.
In some cases the reactions were direct, they turned away at once
without preliminary movements. Specimens with both eyes re-
moved do not react exactly as normal individuals, but they do
move in general away from light. The rate of locomotion in these
is not appreciably affected, but the removal of the anterior end
greatly retards the rate of locomotion. Specimens with one eye
removed orient themselves accurately to light when illuminated
on the normal side, but do not when stimulated in this way on
the blind side.
According to Taliaferro, light must strike a given rhabdome
oarallel with its longitudinal axis in order to cause stimulation.
'Thus, the position of the longitudinal axis of the rhabdome re-
sults in a localization of photic stimulation." It is ijossible, accord-
ing to this investigator, to ex])lain the localization of photic stim-
ulation in one of two ways. P^irst, the refractive central region of
the retinula acts as a sort of lens to concentrate the light on the
sensitive rhabdome. Second, bv assuming a certain structure of
the rhabdome coupled with a shading action of the pigment-cup.
He.sse, '97, ascrii)es the localization of the stimulus entirely to the
pigment-cup.
TrematoDA. In monogenetic forms such as TristoniKni Lang.
18S1, or Ki>id('Ua. Heath, 1902. Iht- brain consists of a rather short,
semicircular band near the dorsal surface just in front of the
nharynx. From it six longitudinal nerves arise, four ventral and
two dorsal. These extend the length of the body and end in the nn<5-
terior sucker. Many small nerves .spring from the brain and the six
FLAT WORMS
39
longitudinal cords. A short distance from the brain the anterior
nerves are united into a curved ganglion and from this a number of
branches run to the anterior end of the body. On the mid-dorsal
line a small median nerve in Epidella runs towards the head and
towards the sucker, but was not found farther than this.
In the main nerve strands and ganglionic areas bipolar cells
are of frequent occurrence and generally one branch from each
might be traced close to the surface of the body while the other
fiber passes into the brain. In a few cases the fibers pass to the
opposite side of the ganglioa or brain before they terminate. Cells
with three branches in Epidella were found with one process to the
Fiji'. 10. Trematode worms. A, B, D, Monogenetic forms. C, E, F. G, Dige-
netic. A. Nervou.s system Tristomum. B. Head end of Epidella,
Health, D. Eyes of Epidella, Health. C. Amphistomum, Loos. E.
Sensory cells of trematode, Havest. F. Nervous system of Cerca-
rineniim, Bettendorf. G. Nerve plexus Corcoriaentim, Bettendorf.
brain, another to the substance of the sucker of the same side, and
the other crosses over to the sucker of the opposite side.
In Epidella. the large mass of nerve fibers and the more numer-
ous longitudinal bands on the ventral side are explained by the
fact that this side rests against the host.
There are four eyes in Epidella. In other forms they seem not
+0 NERVOUS SYSTEM AND SENSE ORGANS
always as well developed and may not always be functional. In
this form the eyes appear as four small pigment spots partly im-
bedded in the dorsal surface of the brain. In this and in Tris-
tumion, each eye-spot consists of an almost spherical, highly refrac-
tive transparent body which in many cases contains one or two
small vacuoles, but a nucleus was not seen. The lens is partly
covered by a cup of dark bi'own pigment granules. These parts are
imbedded in a rather large ganglion cell. Two or three fibers arise
from each ganglion cell and e.xtend some distance into the brain.
A series of delicate muscles are near the eyes and their contractions
bring about rotations of the eyes. One pair of eyes has been found
to move simultaneously with the other, although this does not always
take place. If the animals are vigorous the movements of the eyes
may take place with the rapidity of a heart beat.
The eyes are situated on the dorsal side of the brain. The
tissue between them and the ventral side is clear and light passing
under the host must strike the lens and affect the retina as the
pigment is placed in the most favorable position in the anterior side
of the lens.
In some digenetic trematodes the nervous system has a rather
complicated system of branching as shown in Amphistumitm by
Loss, 1892. Nerve tracts are clearly defined and nerve cells,
although chiefly centered in the broad brain, are also found out
along the peripheral nerves.
Faust, 1918, has studied the eyes in digenetic trematodes. In
twenty-eight species, seven possess pigmented eyes and four non-
pigmented ones. Binoculate species usually have the eye spots in
direct connection with the po.sterior dorsal nerve trunks. In one at
least connections were with the anterior dorsal rami. The central
eye of trioculate species is fused to the anterior dorsal nerve trunk
by a blunt fiber from below. The eye spots consist of a cluster of
rather dark-brown granules forming a deep cup. Within the cup
is a spherical body barely touching the pigment granules. This is
the enlarged nerve ending with a nucleus within.
The development of the eyes in Ccrcaiia f/iV/o.s is as follows:
A branch of the posterior dor.sal nerve with a single nucleus
pushes out from the nerve center to the dorsal margin of the
embryo. As it reaches a position near the surface, the ectodermal
layer of the embryo pushes inwards ju.st posterior to the nerve, so
that a pocket is formed with the opening opposed to the nerve cell.
The end of the nerve fiber enlarges and twists about the inner wall
of the pocket so that the end with the nucleus comes to lie within
the cup. At fir.st the ectodermal cells are evident, but later they dis-
appear. Pigment granules are not present until the nerve ending
comes to occupy its position within the jwcket. (lolden-brown pig-
ment granules come to be formed between the nerve endings and the
FLAT WORMS
41
ectodermal cup. The cell within the cup enlarges and becomes the
lens. The lens is in this way derived from the nerve center.
In Cercaiiaenum Bettendorf, 1897, shows six longitudinal
strands from the brain, with many branches to the pharynx and the
suckers. A complex nerve plexus of nerve fibers and nerve cells is
found over much of the body. Especially are bipolar sense cells
found in the pharynx. Similar bipolar sense cells are demonstrated
by Havet, 1900, by the Golgi method.
Cestoda. The scolex contains the greatest concentration of
the nervous system although in Gnjocotyle there is fully as great a
Fig. 12. The sketch at the top is from a section across a young; flatworm
showing the brain as a dark mass in the left side. The figure at the
left below is from a larval flatworm showing the position of twelve
simple eyes. The middle and lower left hand figures are from em-
bryonic stages of a nemertinian worm showing the developing nervous
system on the left and shown darker in the figures. Salensky.
mass of central nervous system in the caudal end of the animal.
The suckers or other appendages of the scolex region are supplied
with special branches. In some forms there is a definite ring of
fibers. In all two larger and usually four smaller longitudinal
strands run the length of the animal.
Blanchard, 1847, dissected the nervous system in Ligula where
he found a mass of nervous tissue in the scolex with strands run-
+2 NERVOUS SYSTEM AND SENSE ORGANS
ning through the body, especially two thick ones. Moniez, 1881,
found the commissui'es in the forward end of the body.
Lang, 1879-82, figures and describes the nervous system of a
member of the Cestoda where he finds a concentration in the scolex
region and nerves running from this center to the appendages in
this region when present and also long nerves which run the length
of the body.
Roboz, 1882, shows the central ganglion and an extensive nerve
network in cestodes. Some authors claim to have seen ganglion cells
along the nerve strands and in fact Kahne considers the chief longi-
tudinal strands as central organs.
Haman, 1885, also describes the long nerve fibers as having
ganglion cells on them.
Niemeic, 1886, in Li(/ula shows a central ganglionic mass with
two thick strands leading from it and four or more smaller ones,
some of which branch again. Blanchard found similar conditions.
In Schlistvcephaliis, Moniez gives a brief description of the
nervous system also Niemiee, 1886.
In BvtliriuceijIialKK. Niemiee gives some indications of commis-
sures in the scolex region.
In Taenia, Blanchard gives some indication and Moniez dis-
tinguishes a nerve ring in the tip of the scolex. Blumberg, 1877,
finds a larger number of longitudinal nerves than the last author
and Nitsche finds ten strands from the neck region of Taenia.
Niemiee, 1886, finds a nerve ring in the rostellum and eight
nerves coming from the ring. As each one leaves there is a
swelling on the ring with small ganglion cells. A commissure sur-
rounds the central ganglion. Other commissures were also found
in this region.
In ArantliDhi.tln iiivi I'intner. 1881. was one of the fir.st to de-
.scribe the nervous system. Niemiee shows it with branches to the
forward region, a ring commissure below the main ganglion and
with two thick and other thinner longitudinal .strands.
In Tetrarlnpirlnis Lang. '82, was one of the early students.
Figure 11-L, is from another species which resembles the condition
in Tetrarliynclius.
The nerve cells of Cestoda differ greatly in size. Niemiee gives
figures from the cells and nuclei of a number of species. He finds
them to be from 12x16 microns to 28x34 microns cell body;
nucleus, 5x8 microns to 9x13 microns.
Among the more recent literature is the work of Tower, 1900,
on M'Diiezia. The complicated nervous system of this species is
shown in Fig. 11, A.
FLAT WORMS
43
Kofoid and Watson. 1910, call attention to the similarity of the
nervous structures in the scolex of cestodes with that of the pos-
terior region of some trematodes, and they suggest that with Gyro-
cotijle as an intermediate type the scolex part of the nervous system
of tape worms represents the caudal end of the worm.
The only sense oi'gans of tape worms are represented by very
simple end knobs of sense cells in the cuticle. Fig. 11, B.
Fig:. 11. Nervous system cestoda. A. Moniezia, Tower. B. Sensory cells
ending in hypodermis, Zernecke. C. Nervous system Gyrocotyle,
Kofoid and Watson. D, E, F, G, H, I, J, K. Central nervous systems
scolex end several species of Cestodes. L. Rhynchobothrium, Lang.
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FLAT WORMS 45
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FLAT WORMS 47
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48 NER\()LS SYSTEM AND SENSE ORGANS
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'^ NOV 17 1939 i^'^
VOLUME THIRTEEN NUMBEK THREE
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
SEPTEMBER, 1921
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT 0/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
A List of California Arachnida
V. Phalangida, L. Myers J9
IV. ACARINA, F. Cox. p. Jaliraus, IV. Moore - - - 23
Nervous System and Sense Organs
VI. W. A. Hilton 49
Journal of Entomology and Zoology
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A List of California Arachnida
V. PHALANGIDA OR HARVEST MEN
L. Myers
First three figures from Banks.
CosMETlDAE. Second pair of legs without endiles. Peciiplalps shorter than the
body. Eye tubercles low.
Cynorta liimaculata Bks. San Diego. No spines or tubercles at caudal end of
the body.
Phai.angodidae. Hind coxae united to first abdominal at base, free at apex.
Second pair of legs distinct endites. Pediplalps large. Spiracles indistinct.
Sitalces californicus Bks. Martin Co. and Mt. Shasta.
Sehrohunus rohuslus Pack. Mt. Shasta region.
Srotolemon cnlifornica Bks. Alabaster Cave, Calif.
Phai.anciidae. Last segment of the pedipalps long and armed with a claw. Coxa
of fourth leg is united near its base on the posterior side to the tracheal sternite of
the abdomen. Tibial spiracles are present.
Protolop/ius tuberculatus Bks. Gray to brown, more or less mottled. Abdomen
often red-brown. Claremont, Santa Catalina, Santa Rosa.
P. singularis Bks. Near San Diego.
Mitopus lalifornliiis Bks. Los Angeles. CJray above, mottled, femora and tibia
brown.
Gluhlpcs spinulatiis Bks. Red-brnwri, base of legs yellowish. Eye tubercle low.
S. Calif.
Leptobruniis laliforn'nus Bks. Whitish above, mottled with brown and black.
Indefinite vase mark. Los Angeles and S. Calif.
Euryhunus bninneiis Bks. Body very smooth; fourth leg nearly as long as sec-
ond. S. Calif.
E. spitiosus Bks. Grav abo\'e, black mark on each side of base of abdomen.
Femora I and III brown, with a pale ring on middle.
20
Journal of Entoniolony and Zoolog)'
I.eiohumim liimiuulatuin Hk>. Dark lirown, twd prominent yellow >pots. Near
San Diego.
L. fxilififs Wood. Female dark ro>e mark on dor>al >idc. I'rom N. Calif, to
Claremont. * ommon in mts. near Claremont.
ISCHVROPSAI.ID.VE. Last segment of pedipalps shorter than next to last, without
claw. Coxa of fourth leg not fused with adjacent slernite of abdomen. No liliial
spiracles.
Turin III sf'iniisiij Bks. I'alc yellow, claw of mandihies rcd-hrowii. S. t'alif.
T. fallipei Hks. Mt. Shasta.
Nem.\stom.\tid.ae. Stermltes of ainlomeii free, overlapping, and without median
divisional sulcus. The first and second abdominal slermites narrowed in front and
extended betwen coxae.
SemaslDma mojrsla Bks. Back brown to red-browii. Legs pale. From eye
tubercle backwards a row of tubercles, flat tops broader than base. Mt. Shasta,
Claremont.
Troculidak. Slermites of abdometi except genital and anal, fused, do not over-
lap. They have a median longitudinal sulcus. The first and second abdominal
sternites widely rounded in front and overlap the proximal parts of the two posterior
pairs of coxae.
Orlholasmn /<iitififi Bks. Kyc tubercle. Four to five openings on a side. Huni-
b<dl Co. and Ml. Wilson.
(). rugnsa Bks. Common in S. Calif.
Pomona College, Claremont, California
21
Dendidlasma mirabilis Bks. Coulterville, ("alif.
Pomona Jour. Ent. 191), p. 412. Hull. Ill Nat. Hist. 1889 N. 3, p. 99.
Figurf ahi>ve, l.rinhunum lnmiKulaliim. Below, I'rnlolophus tuhtr(utatu3. Figure
ai the right, body of Ortholaimn /'iflif'rs.
A List of California Arachnida
VI. ACARINA OR THE MITES AND TICKS
F. Cox, P. Jahraus, \V. Moore
Figures from Hall, except the plate.
EUPODIDAE. Body divided into cephalothorax and abdomen. Palpi without thumb.
Beak small. Eyes when present near posterior edge of the cephalothorax. Body soft.
Moderate to very long legs. Palpi short. Mandibles small but chelate. Mostly on
ground, predaceous.
Eupodes brevipes Bks. Body red, legs clear. Slender. Sides concave. Laguna
Beach.
Rhagidia pallida Bks. Under stones, Claremont.
Penthaleus hicolor Bks. Spherical, dark body, red legs. Common Claremont.
Bdellidae. Snout mites. Skin not hard. Palpi 4-5 segments. Cephalothorax
large, well separated from abdomen. Palpi large geniculate and bearing long tac-
tile bristles. Mandibles chelate. Body elongate. Lives in moss, dead laves, etc.
Predaceous.
Bdella peregrina Bks. Claremont, Chino.
B. lata Evving. On live-oak, under stones, etc. Claremont.
B. californica Bks. Body white, legs, palpi yellowish beyond base. Body nar-
rowed in front to beak. Eye each side cephalothorax, four hairs in front, longer one
each side beyond eye. Abdomen a few short hairs above. Legs rather slender.
Clarmont.
B. utilis Bks. from black scale.
Anystidae. Coxae contiguous, radiate. Legs slender, bristly. Body few hairs.
No dorsal grove. Tarsi not swollen.
Erythraeus posticatus Bks. Palpi slender, a long thumb. Body dark red, legs
pale. From bark of eucalyptus, Claremont.
E. augustipes Bks. Under stones, Claremont.
E. hiltoni Bks. Claremont.
Erythaeus sp. not mature, on phalangid, Palmer's canyon near Claremont and on
horned toad Laguna Beach.
Tarsotomus terminalis Bks. Body slightly constricted in middle. Two eye spots
in cephalothorax. Many long erect bristles. Claremont.
T. macrnpalpis Bks. Large species sparce bristles, body nearly twice as long as
broad. Claremont.
Tetranychidae "Red spiders". "Palpus with thumb, body well clothed with
hairs. Legs I and II without spine-like processes. Coxae not radiate. Legs usually
in groups of two each. No dorsal grove on cephalothorax. Tarsi not swollen. Man-
dibles for piercing. Hair on body usually in four longitudinal rows. Body oval, few
bristles. Suture between second and third pair of legs. Red, two to four eyes. Pedi-
palps four jointed, usually a strong claw on next to last joint.
24 Journal of Entomology and Zoology
Tflranyc/ius sirn/'lex Hks. Date palm, El Centro.
T. mylilaspiJis Riley. S. California on orange. Tliis i> ilic "citrds rc^l-^pil^cr".
Ked in color, bristles arise from tubercles.
7". sexmuiitlalus Riley. In San Diego Co. in colonies in depressions covered
with silk.
T. himmulalui Harvey. Dn friiil trees, and food plants. Common on man)
plants.
Tflranyclioiiies californiius Bks. l)n citrus trees.
Trnuifall>us itilifornii iii Bks. Small flat, sometimes on citrus trees, l.iiile dam-
age.
Caliyoniis lirminalis Hks. Reil body. Cliula Vista, San Diego. On lemon leaves,
not abundant or important.
Bryobia fralrmis Garman. In Kast called clover mite. In Calif, called almond
mite. S. Calif, and north. Long front legs, four scale-like projections on frimi
margin.
Rhvchoi.ophoridae. Skin not horny. Ceplialothorax without special hairs. Legs
in two groups. Palpi with last segment a thumb, while next to last ends in a claw.
Cephalothorax large on same plane with abdomen, dorsal groDve present.
Rliyneholoplius moeslus Bks. Red. Monrovia.
R. arfniinlii Mali. Bright red or straw color. Dr> sand l.agiina Beach.
R. gratilipfs Bks. Santa Rosa I.
TRO.MniDllD.\E. Harvest mites. Palpi geniculate, ending in one or two claws and
with a thumb at the end. Coxae in groups. Body thickly doited with short hairs,
larsi often swollen. I'ephalothorax small and almost completely hidden by the pro-
jection of the anterior pan of the abdomen. Mandibles for biting. Body globular
Pomona College, Claremont, California
25
or elongate, red, hairy, usually transverse suture between seiond and third Igs. Eyes
often stalked. Legs with two claws. Larva three pairs of legs. Parasitis on spiders,
flies, etc.
Trombidium perscabrum Bis. Red, length 1.4 mm. Peculiar knobbed hairs.
Claremont, also fresh-water pool Laguna Beach.
T. claremonii Bks. Evey's canyon near Claremont.
T. parificum Bks. Dark red. From ants' nests, and from Evey's canyon.
Trombidium sp. Near Camp Baldy.
HvDRACHNiD.^E. Fresh-water mites. Mouth-parts not in a beak. Usually suckers
near genital openings. One or two pairs of eyes. Body oval or spherical, some-
times of large size, often bright colored. Legs usually five-jointed with swimming
hairs. Often attached to aquatic insects.
Hydrachnid, Larvae on notonectid, Claremont, on carabid beetle Laguna Beach.
Hydracna sp. "Probably new" Banks. Large dark red-brown, spherical, found
in great abundance at Laguna Lakes July and August, 1915.
H.^LACARIDAE Salt-water mites. Body rather elongate. Usually a suture between
the second pair of legs. Rostrum often large. Usually three eyes. No swimming
hairs on legs. Mouth in a distance back, no ventral suckers. Lives upon algae.
Pntilacaraiis (atifoniinis Hall. Under stones low tide.
26
Journal of Entomolony and Zoology
Poularai lintt nuiiala Hall. Boih liijiliK arclitd jjlolnilar. I.agiina Bcacli tide
pool.
(.ii/<i,lufinalliiis (iirliij Mall. Tiilc pool I.aguna Beach.
Pomona College, Clareniont, California
27
C. ealiforniius Hall. Tide pool Laguna Beach.
G.\viASlD.\E. Scavenger mites, body broad, short legs, no eyes. Mandibles usually
chelate. Pedipalps five-jointed, legs six-jointed ending in two claws. First pair of
legs inserted at one side of the mouth opening. Male genital opening usually on
anterior margin of sternal plate.
Gamasus calijornicus Bks. Body yellowish, legs paler.
Parasitus frontalis Bks. From wild mouse, Laguna Beach.
Parasitus sp. Free living, Claremont, Chino.
Macroihflfs sp. Chino swamp.
28
Journal of Entomology and Zoology
Seius orchesloidear Hall. Female light straw color. Male lighter. Dorsal plate
over whole back. Ovoid. From the amphipod UrchtsloiJra eaiiforniana, Laguna
Beach.
Laflafs pilosiilii Bks. Santa Rosa I.
Uropodidae. With a distinct spiracle on lateral sligmal plate abo\-e 3-4 coxae.
First pair of legs inserted in same opening as mouth-parts. Back of body extending
towards and hiding mouth-parts from above.
Pomona College, Clareiiiont. Calitoinia
29
Uropoda sp. ^'oiing on caiahici beetle and on Stnlnpendrii.
Dermanvssidae. Mandibles for piercing. Body sometimes constricted. Para-
sitic on vertebrates.
Dermanyssiis gallinae Redi. Parasitic on chickens.
Orb.atidae. Horny beetle mites. Cephalothorax with a special hair on the pos-
terior lateral vertex. Skin hard. Abdomen wtih wing-like expansions. Body minute,
divided into two parts by transverse suture. Mouth-parts small hidden. Live upon
vegetable or decaying material. Palpi five-jointed.
Hfrmannia hirrngtyphua Hall. Brown, black markings, mandible chelate. Roug
deep sculpturing. Claremom.
30
Journal of Kntomology and Zoolog)'
Orihata liumiJa Hall. • olor rhcsliuil, polished. Abdomen «ilh wings. Mandible
chelate. Laguna Beach under board.
(). californiia Bks. Abdomen red-brown, basal joints of legs brown, rest yellow-
ish-broivn. Cephalothorax brown. Ml. Shasta.
(). alula var. laiijorniia Hall. Black, polished abdomen with wings. Claremonl.
Pomona College, Claremont, California
31
Phthiracarus cryptopus Bks. Body brown, yellowish at base of abdomen. Smooth
shiny, legs pale. Cephalothorax six bristles above, anterior pair shorter than others.
Abdomen large high, about one-fourth longer than broad, two rows of tine hairs
each side above. Legs very short and hairy. Claremont.
Ertmaeus bilamellaius Hall. Claremont under leaves.
E. modestus Bks. Trunk and branches orange trees. Live upon plant life grow-
ing on trees.
Sotaspis pectinata Hall. Yellow brown, smooth, polished. Claremont, Calif.
iV. bilnmellatus Hall. Light chestnut, smooth not polished, without wings. Man-
dibles large chelate. Follows Michael, near A', burrnivsi, but differs in having no
hairs on abdomen. Under stones Claremont.
A', nuda Hall. Black, smooth polished. Mandibles chelate. Under boards,
Claremont.
Journal of Kntoniolony and Zoology
I'aralioJes intiimita Hall. Dark lirowii, almost black, stout chelate.
I.ii/imiinniii sf'inojd Hall. Legs colorless, skin clear. Maiulililes lieav> chelate.
I.iaiorus mnjfslus Bks. Body pale, red-hrown, le^s pale yellow, fephaloihorax
four ridges, and four bristles above.
Kriophvidaf. (iail miles. Body small, wurmlikc caudal end elongate. No eyes.
Two pairs of legs. (Jails always open.
Pomona College, ClareniDnt, California
33
Paraphytoptus californiius Hall. (Possibly may be P. peravorus.) Gall on
Artemisia. Abdomen anulate.
Eriophyes oleivorus Ash. Silver mite.
Tarsone.\iid.\e. No ventral suckers. Legs end in claws, body divided into cep-
halothorax and abdomen. Female with clavate hairs between legs one and two.
Tarsonemus approximatus Bks. Pomona, Calif. Under Cilricold scale.
T. assimilis Bks. From red scale. Whittier.
Tyroclyphidae. Small, elongate, smooth. Legs alike. Chelate mandibles, no
eyes. Palpi close against mouth parts. Legs long, clavate hair on tarsi of one and
two. Not parasitic except a few on bees. Mostly live on organic matter. Cheese
mites, etc.
Tyroglyphus longior Gervais. Hairy bristles on body, long tarsi. Calif.
T. americanus Bks. From lemons in storage S. Calif.
Tricholarsus xylocopae Donn. European species found on Xylocopa californica.
34
Journal nt KntnmDlog) and Zoology
Rhizoglyt'lius loni/istrialiu var. (nlijnrniius Hall. From Kalinin^, injury to bark
of apple tree.
R. Inisalis Bl»>, SprcikcU. Calif., nil sii^ar lieet.
R. rhizBf>liaiius Bks. On onions, Calif.
Olyriphn'/iij nliniis Bkv. Berkeley, Calif.
Car/ioglyp/iiis fiassutai urn llering. From Fresno on dry ti({s.
Pomona College, Claremont, California 35
ASAI.GESTIDAE Bird mites. Small, elongate, transverse striations on the body.
rierotiyssus hifurmlus Hall. Integument strongly cliitinized, from PetfiodieliJuii
hinifroru.
36 Journal of Entomology and Zoology
THE TICKS
Arcesidae. No dorsal shield, head hidden under front of body. Skin rough
coxae usually contiguous or nearly so. Tarsi without apical pulvillum.
Argat miniatus Koch. Riverside.
Ornithodoros coriaceus Koch. San Francisco and Santa Clara Co.
O. megnini Dug. Red brown to black. Los Angeles.
U. lalaje Guer. San Clemente Island.
IXODIDAE. Back covered by a horny shield, head distinct from the body. Anus
in middle of ventral side. Skin finely striated. Tarsi with pulvillum. Male almost
entirely covered with dorsal shield. Female shield only on anterior part of dorsum.
Ixodes hexagonus. Santa Clara Co., Mt. Shasta.
/. californUus Bks. Laguna Heach, Claremont, Sania Clara Co. On fox and
deer, dog. Shield red-brown, paler in middle, body brownish or yellowish, cojtae
brown, legs paler. Few hairs. Shield long, finely punctured.
/. angustus Neum. Siskiyou Co.
/. seulptus Neum. Santa Cruz Mts., Calif.
/. fralti Bks. Claremont.
Irgas miniatus Koch. Large ticks, exact location of capture not known. Calif.
Ornithodoros megnini Duges. Mt. Shasta ; also S. Calif.
Dermacentor occidentalis Neum. Mis. near Claremont and foothills.
I), reticulatus Feb. Palo Alto and Mt. Shasta.
I), paruma/ierlus Neum. Lake Side, Calif.
O. occidentalis Neum. Santa Clara Co., Humboldt Co. From deer.
Ceralixodes signatus Birula. Cormorant, Pacific Grove.
.Imhiyomma maculatum Koch. Tulare Co., Calif.
./. cajennense Beb. San Diego.
Ilaemafiysalis lef>oris-plaiistris Pack. Dn rabbit, Claremont.
//. concinna Koch. Claremont, on rabbit.
Jour. Ent. Zool. VI, 1914, pp. 56-60. VIII, 1916, p. \2. Trans. Am. F.nl. Soc.
XXI, 1894, p. 22. Proc. Calif. Ac. Sc. Zool. MI, 1904, pp. 365-369. Hubbards
Orange G. Insects 1885, p. 216. Jour. N. Y. Ent. Soc. 1904, pp. 54, 55. 1st Laguna
Report. Pomona Jour. Ent. II, p. 280, III, p. 510. V. S. Dep. Agr. Tech. ser. 13,
1906, pp. 12, 20. Trans. Lin. Soc. XI, 1815, p. 397. Mem. Soc. Zool. Fr. 1899, p. 136.
Arch. f. Naturges. X, 1844, pp. 219, 237. La Natur Mex. VI, 1883, p. 196. Ent. Sysi.
IV, 1874, p. 428. Banks, Tyroglyphidae, V. S. Dep. -Agr. Tech. ser. 13, 1906. Banks,
lodoidea, l". S. Dep. Agr. Tech. ser. 15, 1908. Banks, .Xcarina V. S. Nat. Mus. 1904.
Quayle, Red spiders and miles of citrus trees. Bull. 234, Berkeley, 1912.
Pomona College, Clareiiiont, California
37
IxoDlDAE A. Haemafi/iysalis iepnru-paluslris, fresh and gorged female. TvROG-
I.VPHIDAE B. C.arpoglyplnts passularum, C. Gtyciphagus ohesits. Eriophyidae E. Erio-
phyes oleivnrus. Tetranvchidae D. Telranychus sexmaciilatits. F. Tenuipalpus cali-
fornicus, G. Telranyehoides lalifoniims. H. Caligonus terminnlis. J. Tetranychus bi-
macutalus, K. Bryohia praletuis. 1.. Telranyilius mytilaspidis. Orbatidae 1. Eremaeus
moilrslus. TvRnGl.VPUIDAE M. Tyrnglyphus aineruauus.
VI. Nemertinea
The first work of any importance which deals with the nervous
system of these worms is that of De Quatrefages in 1846. He de-
scribes the central nervous system as composed of two distinct
lateral lobes united below and above by commissures. From the
lateral lobes two more or less isolated longitudinal bands extend
themselves towards the posterior end of the animal. So far as the
figures are concerned this early work is even more detailed than
that of M'Intosh in 1873. The more recent information about this
interesting group has been furnished especially by Hubrecht in
numerous papers from 1875 to 1887. Although the cellular details
are not shown, the relative position of the central fibrous core is
given in relation to the surrounding nerve cells. He also clearly
distinguishes the dorsal median nerve springing from the slender
dorsal commissure. The dorsal and ventral lobes of the brain are
shown more clearly than in earlier writings. In Eupolia a dorsal,
middle and ventral lobe are shown.
Hubrecht in his two papers of 1887 suggests the neniertineans
as a group of animals valuable in tracing the relationship of the
vertebrates and invertebrates. He bases his hypothesis largely
upon the arrangement of the parts of the nervous system. In the
group there is some variation in the extent and position of the
lateral nerve cords and in some, the mouth opens behind the brain
and in some in front of the brain. Such facts as these give sug-
gestions of an intermediate condition between annelids and arthro-
Dods on the one side and vertebrates on the other. Other writers
have compared the large lateral nerves of nemertineans with the
central nerve cords of .some I'ound worms.
Biirger in a number of works from 1883 to 1895, has made a
considerable studv of the nervous system by various methods. He
has also studied the histological details of the nervous system. His
papers are the most comnrehensive and important in this field.
Biirger de.scribes the nerve cells as all unipolar and uninclosed in
soecial membranes. He classifies nerve cells as follows: (1) The
smallest cells sensory in nature; (2) medium sized cells; (3) large
cells: (4) very large cells, the so-called "Neurocorde" cells.
Montgomery, 1897, agrees with Burger in many respects, such
as uninolar condition of the nerve fibers, but these are composed
of "a homogeneous un.staining axis cylinder which is probably fluid
and a fine spongioplasmic layer."
In Cerehratiihis, the large nerve fibers diff'er from the others in
size. They do not give off" collaterals but divide dichotomously and
are arranged segmentally. The largest ganglion cells are present
in three pairs in the ventral brain lobes and are distributed irreg-
ularly along the lateral cords, but are absent in both ends. In the
50
NKRNOUS SYSTEM AM) SENSK ()R(]ANS
lateral cords they increase in number posteriorly and are more
abundant on the dorsal side. In each lateral cord both dorsally and
ventrally are radial clusters of medium sized cells showing a bi-
lateral arrangement.
Haller. 1889, shows a neuroglia network in Ceiehratiilns and
an anastomosis between the branches of multipolar ganglion cells.
The nemertineans are divided into groilps somewhat by the
position of the nervous system in relation to the body-wall. The
more primitive condition seems to be when the brain and chief
branches are outside the muscle layers, in the epithelium or below
the basement membrane. In some the nervous system is found
in the muscle layers of the body-wall and in others the brain and
chief nerves lie in the parenchyma internal to the mu.scle layers.
Nervous system an'h sense okuans of Nemeutinea. A. Nervou."!
system of Cerebral ulii.i showing chief nerves and the position of the
cential fibrous mass, Hubrecht. B. Set'tion of eye of Drcpanophoriis.
Hubreeht. C. Diapram of head end of Cerebratiiliis. D. Section of
eye of Linens, Pitmiett. E. Brain of DrapaiioplwrH.i, Hubrecht. V.
Briin of Riqitilia. showing fibrous core on the risht. Hubrecht. G. H.
Cross sections through brain of RitpoUa, left side and oesophajrus
shown in each. I, J. Scheme of some nerve cells and fibers in the
lateral cord and ventral panplion in Annpla, and Drepanophoms.
Biirjrer.
Hul)recht, '87, suggests that the more primitive nervous system
of these animals has a most complicated intricate network of peri-
NEMERTINEA
51
pheral nerve tissue. This network suggests the "most ancient
arrangement of the nervous tissue." In the more highly specialized
forms, the brain and lateral nerves are more concentrated. Prob-
ably all nemertineans have more or less peripheral nerve networks
even though Hubrecht might not have seen them by his methods,
but the fact remains that those forms in which the network is
especially marked are more primitive because of it. Montgomery
believes that Haller is mistaken as to the multipolar condition of
these cells.
Fig. 14. Reconstruction of the nervous system of Carinella shown from the
ventral side. Figure at the left, side view of a reconstruction of the
upper portion of the central nervous system of Carinella. The figures
at the right are from cross section taken at various levels. The upper
and the two lower figures are from one side only. X75, Hilton.
In general the central nervous system of the Nemertinea is as
follows : A brain composed of two ganglionic masses at the anterior
end of the body, on on each side of the proboscis. These are united
52 NER\()US SYSTEM AND SENSE ORGANS
by ventral and dorsal commissures passing about the proboscis.
The dorsal band is often more slender than the ventral and from it
a slender dorsal nerve runs the length of the body. Each lateral
brain lobe is often partly divided into a dorsal and ventral lobe.
From each lateral ganglion a large nerve trunk passes back and may
unite with its fellow of the opixisite side just above the anus.
Nerves are given off from the brain to the eyes when present,
and to anterior portions of the body. Two branches come off from
the dor.sal commissure and run to the proboscis. The so-called
vagus nerves arise from the internal borders of the brain not far
from the origin of the lateral cords. They are sometimes united
by a commissure and then pass down the oesophagus.
Eyes are usually present along the sides of the head, sometimes
a single pair, at other times one or more groups on each side. The
eyes in their simplest conditions are mere pigment spotis. in others
there is a clear area filled with fluid which is supported by strands
from cells and held by a limiting membrane. Sensory cells are con-
nected with the brain bj' fibers and with pigment at the outer side.
The .sensory area seems to be like rods in certain forms.
In some cases otocysts have been found on the surface of the
brain. At the anterior tip of the head groups of cells bear long
bristles. In some, these areas are retractile. Taste has been sug-
ge.sted as the function of these "frontal" organs. The .so-called
"side" organs occur as a pair of epithelial patches on each side of
the body in the region of the e.xcretory pore. These have an abun-
dant nerve supply but their function is unknown.
In most forms a peculiar pair of organs is found in the head
region in close connection wtih the brain. Hubrecht suggests that
they may be respiratory. Biirger thought that they might be organs
used for determining the condition of the water. They may be
shallow depressions, longitudinal or slit-like or the slit may be at
right angles to the body. In some, ciliated ducts pass inwards and
penetrate into special lobes called the cerebral organs.
Thompson, 1908, in CerehratuUis laeteus finds six ventral com-
missures from the ventral lobes of the brain. Some of these come
from the fibrous core and some come from the cellular sheath of
the brain. Other commissures are found beyond the brain.
Six pairs of "neurocord" cells and one unpaired cell are found
in the ventral lobes of the brain. There is probably individual
variation as to their number.
The brain is complex but resembles in its form and commis-
sures that of the tubularian worms.
Coe and Ball, 1920, in Nectnnemertes, find both dors^al and
ventral commissures well developed. Cerebral and frontal organs
are lacking.
NEMERTINEA 53
In the blastula of Cerebratulus cells on the apex of the larvae
develop cilia and sink below the general surface. This forms the
apical sense organ of the larva.
The brain of the adult develops by thickenings of the apical
discs.
BIBLIOGRAPHY
Bohming, L.
1898. Beitrage zur Anatomie und Histologie der Nemertinen. Zeit. f.
wiss. Zool. Bd. 64, pp. 478-564, Taf. 13-17. 1 text fig.
Burger, O.
1890. Beitrage zur Kenntnis des Nerversystems der Nemertinen.
Inaug. Diss. Gottingen, pp. 1-76. 4 text figs.
1890. Untersuchungen uber die anatomie und Histologie der Nemer-
tinen nebst Beitragen zur systematik. Zeit. f. wiss. Zool. Bd. 50,
p. 1277, Taf. 1-10. 12 text figs.
1891. Zur Kenntnis des Nervensystem der Wirbellosen. Neue unter-
suchungen uber das Nervensystem der Nemertinen. Mitt. Zool.
St. Neap. Bd. 10, pp. 206-254, Taf. 14-15.
1895. Die Nemertinen des Golfs Neapel. F. und Flora Golf. Neap.
Bd. 22.
1897-99. Nemertini. Bronn's Tier-Reichs. Bd. 4, pp. 1-542, pi. 1-22.
Text figs. 1-43.
Coe, W. R.
1905. On the anatomy of a species of Nemertinean (Cerebratulus lacteus
Ver). Trans. Conn, ac, vol. 9, pp. 480-514, pi. 10-15.
1905. Nemerteans of the west and northwest coast of North America.
Bull. Mus. Comp. Zool., Harvard, vol. xlvii, pp. 1-318, 25 pi.
Coe, W. R., and Ball, S. C.
1920. The pelagic nemertean Nectonemertes. Jour. Morph., vol. 34,
pp. 457-485, 5 pi.
Dewoletsky, R.
1880. Zur Anatomie der Nemertinen. Zool. anz.
1888. Das Seitenorgan der Nemertinen. Arbeit a. d. Zool. Inst. d.
univ. Wien. Bd. 7.
De Quatrefages
1846. Etudes sur les types inferiurs. Memoire sur la famille des
Nemertines. Ann. des. sc. nat. ze ser. Zool. T. 6, pp. 173-303, pi.
8-14.
Haller, B.
1889. Beitrage zur Kenntnis der textur des Central Nervensvstem
Hoher Wurmer. Arbeit Zool. Inst. Wien. Bd. 8, Heft. 2, pp.
1-138. 5 pi. 4 wood cuts.
Hilton, W. A.
1917. A reconstruction of the Nervous System of a Nemertinean Worm.
Jour. Ent. and Zool., no. 3, pp. 119-124. 2 figs.
54 NERVOUS SYSTEM AND SENSE C)R(;ANS
t
Hubrecht, A. A.
1875. Some remarks on the minute anatomy of Mediterranean Nemer-
tinens. Q. Jour. Mic. Sc, vol. 15, pp. 249-257, pi. 13, figs. 6-8.
1880. Zur anatomy and Physiologia des Nervensystems der Nemer-
tinen. Naturk. Verh. der Kominkl. Akad. Decl. XX, pp. 1-40,
pi. 1-4.
1880. Recherehes on the nervous system of Nemertines. Q. Jour. Mic.
Sc. n. s.. vol. 20, pi. 23.
1880. The Peripheral nervous system of the Palaeo and Schizonemertea,
one of the layers of the body-wall. Q. Jour. Mic. Sc, vol. 20.
1881. Studien zur Phylogenie des Nervensystems Nat. Verh. Der.
Konink. Akad. Dael. XXII, pp. 1-19. pi. 12.
Report on the Nemertea collected by H. M. S. Challenger during
the years 1873-1876. Rep. Vovage. H. M. S. Challenger Zool.,
vol. 19, pp. 1-146. N. Syst., pp. 73-90.
1887. The relation of the Nemertea to the vertobrata. Q. Jour. Mic.
Sc, vol. 27.
Ikeda> I.
1913. A new fresh water Nemertine from Japan, Stichostemma grandis.
Annot. Jap., Tokyo, Zool. soc, vol. 8, pp. 239-256, pi. 4.
Kennel, J. V.
1877. Geitrage zur Kenntnis der Nemertinen. .\rbeit. a. d. Zool. Inst.
Wurzburg. IV.
M'Intosh, W. C.
1874. A Monograph of British Annelida. Rav. Soc. Part. I. The Nemer-
teans, pp. 97 213, pi. 11 23. 14 text figs. N. Syst., 81-84.
1876. On the Central nervous system, the cephalic sacs and other points
in the anatomy of Lineidae. Jour. Anat. Phys., vol. 10.
Montgomery, T. H.
1897. Studies on the elements of the Central Nervous System of
Heteronemertini. Jour. Morph., vol. 13, no. 3, pp. 381-444, pi.
24-26.
Punnett, R. C.
1901. Lineus. Mem. Marine Biol. Mem. I.. M. B. C. Mem., pp. 1-37,
pi. 1-4.
Semon, R.
1898. Zoologische Forschungsreisen in Australien. Bd. V, 1\' Lief.
Neue Nemertinen aus Amboina, pp. 593-614, Taf. 47-51.
Thompson, C. B.
1908. The Commissures and the Neurocord Cells of the Brain of Cere-
bratulus lacteus. Jour. Comp. Neurol and Psvch., vol. 18, no. 6,
pp. 641-661. 13 figs.
l^
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
DECEMBER, 1921
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT 0/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
A List op California Arachnida
VII. Araneida, M. Moles, J. Johnson . . - 39
Cphinroidea of the West Coast of North America
Arthur S. Campbell 46
Nervous System and Sense Organs
VII. W. A. Hilton 55
Entered Claremont Cal.. Post-Office Oct. 1, IBIO. as second-class matter, under Act of Congress of
March S. 1878
Journal of Entomology and Zoology
EDITED BV POMONA COLLEGE, DEl'AUTMENT OK ZOOLOGY
Subscription $1.00 to domestic, $1.25 to foreign countries.
This journal is especially offered in exchange for zoological
and entomological journals, proceedings, transactions, reports
of societies, museums, laboratories and expeditions.
The pages of the journal are especially open to western ento-
mologists and zoologists. Notes and papers relating to western
and Californian forms and conditions are particularly desired,
but short morphological, systematic or economic studies from
any locality will be considered for publication.
Manuscripts submitted should be tyjiewritteu on one side of
paper about S by 11 inches. Foot notes, tables, explanations of
figures, etc., should be written on separate sheets. Foot notes
and figures should be numbered consecutively throughout. The
desired position of foot notes and figures should be clearly
indicated in the manuscript.
Figures should be drawn so that they may be reproduced as
line cuts so far as possible. An unusually large number of half
tones must be paid for in i)art by the author. Other more
expensive illustrations will be furnished at cost. Figures for
cuts should be made to conform to the size of the page when
reduced, that is, 5 by 71/2 inches or less. The lettering should
be by means of printed numbers and letters pasted on the
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Authors of articles longer than a thousand words will receive
fifty reprints of their publications free of cost. If more than
this are desired, the order should be given with the return of
the proof sheets. Extra copies and special covers or special
paper will be furnished at cost. Authors of short contributions
will receive a few extra copies of the nimiber containing their
articles.
Manuscripts should be seut by express or registered mail.
Address all communications to
The Joi'RNAL OF Entomology and Zoology
William A. Hilton, Editor
Ciaremont, California, U. S. A.
A List of California Arachnida
VII. ARANEIDA OR TRUE SPIDERS
M. Moles, I. Johnson
AvicuLARllDAE. C'helicera project forward and claw moves vertically. Two
pairs of book-lungs. Coxae of pedipalp like the legs, lacks a distinct endite.
Bothriocyrtum californicum Camb. Los Angeles Co., etc. Common trap door
spider.
Eutychides ^'ersicolor Simon. Santa Clara Valley.
Hexura picea Sim. Mariposa Co.
Brachythele longitarsis Sim. Calif.
B. theveneti Mariposa Calif.
Atypodes riversi Camb. Black Mt., Calif.
EuTpelma calijornica Auss. Santa Cruz and south to Claremont.
E. riUyi Mar. Calif.
E. leiogaster Auss. Calif.
E. marxi Simon. Calif.
Hexura julva Chamb. Claremont.
Nemesoides hespera Chamb. Claremont.
Amblyocarenum talpa Bks. Calif.
Aptostichus atomarius Simon. Calif.
A. claihratus Simon.
A. standfordiiinus Ch. P. Smith, San Francisco Co.
Aficularia calijornica Bks. Calif.
Hebestatis thereneti Simon. Calif.
ArypiDAE. Distinguished from the previous family by more complicated palpus
of male. Coxa of pedipalps bears a large conical lobe. They also have a large
endite on the coxa of palpus.
Aliatypus californicus Bks. Santa Clara Valley.
ULBORmAE. Spin orb-webs. Have cribeillum and calamistrum. Dark eyes, lat-
eral ones farther apart than the two pairs of median ones. Posterior metatarsi much
curved and armed below with a series of spines.
Vlhorus calif fjrniciii' Bks. Napa Co. and near Claremont.
DiCTiNlDAE. Cribellum and calamistrum. Anterior median, eyes dark, others
white. Lateral eyes on each side nearly touching. Tarsi of legs three claws. Ir-
regular web.
Amaurobius nefadensis Simon. Northern counties.
A. nigrellus Chamb. Claremont.
A. pictus Simon. San Francisco.
Dictyna sublata Hentz. Lake Tahoe to Claremont.
D. I'olucripes Keyser. Palo ."Mto to Claremont.
D. calcarala Bks. San Pedro.
D. mians Chamb. Claremont.
40 Journal of Entomology .ind Zoology
Dniynina pallida Bks. Mt. Shasta.
Diilyolal/iys laliforniia Bks. Palo Alto.
I'arauximiis Iradalui Chamb. Claremont.
Auximus pallestfiu Chamb. Claremont.
.4. laleicans Chamb. Claremont.
FlLISTATlD.^E. Eyes massed in small group, anterior median eyes dark, rinind,
rest oval or angular, white. Chelicerae small without condyle, chelate.
Filistata hibernalis Hentz. Mill N'allev to Claremont.
Dysderidae. Six eyes. Four spiracles near base of abdomen. A pair uf lung
slits and a pair of tracheal spiracles. Coxae of first pair of legs long and cylin-
drical.
Segeslria paiifica Bks. Mt. Shasta and Claremont.
Sc\TODlDAE. six eyes, one tracheal spiracle. All eyes white. No siiturf between
labium and sternum.
Dit/uflia caniles McCook. San Diego, 1 os .Angeles.
Fleclreurys suprenans Chamb. Claremont.
Leptonetidae. Six eyes, small long legs, suture beiweni labium :ind sternum.
Leploneta californica Bks. Mt. Diablo.
Usoala gracilis Mark. Calif.
Drassidae. Eight eyes in two rows. Two tarsal claws. Four spinnerets widely
separated. Tarsi with bundles of terminal tenent hairs.
DrassiiJfs lalifornica Bks. Sierra Co. and Martin Co.
D. ctles Chamb. Claremont.
Mrgamyrmfiion falifornicum Simun. San Francisco, Claremont.
Drassinflla modrsta Bks. San Francisco and Claremiint.
Gnapliosa raliforaica Bks.
Poecilochrna pacifica Bks. Sierra Cfi., Stanford and Claremont.
P. motilana Em. Claremont.
P. concinna Sim. Calif.
y.elolfi femoralis Bks. Claremont.
X. matulaltis Bks. Claremont.
'A. pacificii} Bks. Santa Rosa I.
Z. laiho Chamb. I'laremont.
7.. irrilans Chamb. Claremont.
'/.. gynrlhus Chamb. Claremont.
'/.. flhnps Chamb. Claremont.
Ilerpyllus augustij Bks. San Pedro.
//. (alijornifus Bks. Lakeside, Calif.
//. ialiJiij Bks. r.os Angeles and Claremont.
//. pins Chamb. Claremont.
Srrgfoluj hifolor Bks. Claremont.
Pomona College, Claremont, California 41
Callilepis insularis Bks. Guadeloupe I., Claremont.
Pholcidae. Very long legs, irregular webs. Tarsi of legs three claws, usually
eight eyes. Group of three eyes on each side.
Pholcus phalangioides Fuessl. Los Angeles, Claremont.
Physocyctus golbosus Tacz.
Psilochorus californiae Chamb.
ZoDARliDAE. Legs nearly equal in site. Internal face of the endites is not fur-
nished wtih serrula, but bears an apical scopula. Rostrum membranous and furnished
above with a band of hairs.
Lutica maculata Marx. Calif.
Theridiidae. Eight eyes. Three tarsal claws, comb on tarsus of fourth pair of
legs. Chelicera no condyle.
Theridion tepidariorum Koch. San Francisco, Claremont.
T. ptacens Keys. Calif.
T. differens Em. Palo Alto, Mt. Shasta.
T. fordum Key. Santa Cruz.
T. calijornicum Bks. Calif.
T. inconstans Curtis. Calif.
T. sexpuncialiim Emerton. Mill \'alley.
T. pictulum Bks. Calif.
Lalrodfctes mactans Fab. North to south, Catalina I.
Dipoena pictipes Bks. Claremont, Calif.
Argyrodes decorus Bks. Calif. '
A. jucundiis Camb. Los Angeles, San Pedro.
Euryopis jttnehris Hentz. San Francisco.
Steatoda (jrandis Bks. Claremont.
Lithyplianles tedus Keyser.
LiNYPHliDAE. Three claws, eight eyes. No comb on tarsus. Organs of stridu-
lation. Dissimilar eyes. No lateral condyle or chelilerae.
Diptocephalus fasciaius Bks. Calif.
Lniyphia arcuata Keyser. San Francisco.
L. digna Keyser. Palto Alto.
L. phrygiana Koch. Palo Alto.
L. rubrofasciata Keyser. Mt. Shasta.
Erigone lalifornlia Bks. N. Calif and Claremont.
Bathyphautes pallidulus Bis. Calif.
Argiopidae. Orb-weavers. Three claws, eight eyes. Tarsi hairs, nn cnmh.
Tetragnaiha extensa Linn. Alameda Co.
T. laboriosa Hentz. N. and S. Calif.
Leucauge hnrtorttm Hentz. Los Angeles.
Argiope Irijasciata Forsk.
A. argentata Fsb. S. Calif.
A. aurantia Lucas.
A. ai'ara Thorell. Calif.
Ordgarius cornigeriis Hentz. Los .Angeles.
Gasteracantha maura McCook. Claremont.
42 Journal of Enlomolog) and Zoolog)-
C. cancrifornis Linn, talif.
G. tetracantha Linn, i'alif.
Mela menarJi Latrelle. Claremoni.
Cyrtophora latijorniensis Keyser.
Cyclosa itdex Cambs. N. Calif.
C. conica Pallas. N. to South.
Euslala anustera var. lumhlea McCook. Calif.
Ztlla californica Bks.
/. x-nolala Clerck. Claremoni.
Melargiol>e Irifasciala Forsk. <'larrmont.
Aranea angulata Clerck. Clareraont.
A. marmorea Clerck. Claremoni.
A. curcurhilina Clerck. Claremoni.
A. carbonaria Koch.
A. miniala Walck. Claremoni.
A. bispinosa Keys. Calif.
A. conchlea McCook. Claremoni.
A. oaxacrnsis Keys. Sitz. Palo Alio lo Los Angeles.
./. JispiHala Hentz. Mill Valley, Mi. Shasla.
./. labyrinl/iea Hentz. Manin Co. lo Claremoni.
A. I. yrinelli Coolidge.
A. nepliiloiJei Camb.
A. Irifoliiim Hentz.
A. palagiala Clark. X. Calif.
A. paiifiin McCook. N. and S. Calif.
A. californiia Bks. Calif.
A. gemma McCook. N. to S.
A. variolala Camb. Calif.
A. gosogana Chamb. Calif, desert region.
Leucauge argyra Walck. Calif.
Ctenid.^e. Wandering spiders, usually. Eyes three to four transverse rous.
Ends of endites clothed in dense uneven hairs. Two-clawed.
Titioliij (atifornicus Simon. From Calif.
Clubionidae. Flat tubular nests, eight eyes in two rows, two tarsal claws. Lower
margin of furrow of cheliccrae distinct, armnl »iili lecili. Tarsi usualK with bundle
of tenent hairs.
Gayfnna lalijornidl Bks. Palo Alio, Mill \allc\.
Chirm nnlliium indusum llenlz. Mill \ allev. Clareinoiu.
Clubionii pacificn Bks. Claremoni.
Olios fascieulalui Simon. Calif.
O. schistiis Chamb. Claremoni.
Anyphafim (rebrispina Chamb. Claremoni.
A. rums Chamb. Claremoni.
.'/. zina Chamb. Claremoni.
A. innirsa Chamb. Claremoni.
A. niinilflla Chamb. Claremoni.
Pomona College, Claremont, California 43
Anachemmis sober Chamb. Claremont.
A. dolichopus Chamb. Claremont.
Namopsilus pletus Chamb. Claremont.
Micaria palliditarsus Bks. S. Calif.
Castaneira descripta Hentz. Claremont.
C. pacifica Bks.
C. tricolor C. Koch.
Trachelas tranquillus Hentz. Claremont Mts.
r. calijornicus Bks. Claremont.
mike trivittata Keys. Calif.
ACELENIDAE. Three claws, usually eight eyes. No scopula on tarsus. Trochan-
ters not notched. Hind spinnerets very long. Funnel-web weavers.
Agelena pacifica Bks. N. Cal., Catalina 1. and Claremont.
A. californica Bks. Stanford, Claremont.
A. naevia Hentz. Claremont and Catalina 1.
A. rua Chamb. Claremont.
Tegenaria domeslica Clerck. Claremont.
T. californica Bis. N. Calif and Claremont.
Cybaeus reticulatus Simon. Claremont.
f,'. minor Bks. Claremont.
C/iorizotnma californica Sim. San Francisco.
Cybaeodcs incerta Bks. Salton, Calif.
Coelotes esaplus Bks. Calif.
MiMETiDAE. Tibia and metatarsi nf first two pairs of legs with very long spines
and shorter between.
Mimetus interfector Hentz. Claremont.
Thomisid.ae. Crab-spiders. First and second pair of legs usually longer than
third and fourth. Eyes small dark, two rows usually recurved. Lower margin of
chelicerae indistinct, unarmed, upper unarmed or with one to two teeth.
Xysticus californicus Keyser. N. to S.
X. formosus Bks. Mt. Shasta.
X. ferox Hentz. Claremont.
X. gluosus Keyser. Claremont.
,\'. triguttatus Keys.
.\'. montanaensis Keys. Calif.
Coriarachne brunneipes Bks. Mt. Shasta.
Runcinia ateatoria Hentz. N. Calif.
Misurnena vatia Clark. N. to S.
Misumesstis pictilis Bks. Palo Alto.
M. pallidulus Bfes. San Francisco.
Misumenoides aleatorius Hentz. Claremont.
M. californicus Bks.
Misumenops asperatiis Hentz. Claremont.
M. californicus Bks.
M. importunus Keys. Calif.
M. diegoi Keys. Calif.
44 Journal of Entomology and Zoology
HI. moJeslus Bks. Calif.
M. munieri Coolidge.
M. pallidulus Bks.
A/, pklilis Bks.
Tmarus magniceps Keys, l.os Angeles.
Thanatus coloradcnsis Keyser. N. and Clarcmont.
T. retenlus Chamb. C'laremont.
T. oblongus Walck. Palo Alto and south.
Phitodromus rujus Wale. N. t'alif.
P. calijornicus Keyser. N. Calif.
P. moestus Bks. Claremont.
P. pernix Blackwall. Claremont.
Lycosid.ae. Wolf-spiders. Trochanters of legs notched. Lorum of two pieces
one notched to receive the other. Eyes in three rows, posterior lateral eyes behind
posterior median, first row of four small eyes, two back rows of two large eyes each.
Lycosa pacifica Bks. N. to Claremont.
/.. brunneiventris Bks. Halo .^Ito, Claremont.
L. koclii Keys. Claremont, and Ontario Mt.
L. ferriculosa Chamb. Claremont.
L. piratimorp/ia Strand. Calif.
L. ramulosa McCook. Calif.
Pardosa slenialis Thorell. Claremont.
/'. lapidicina Em. Claremont.
P. tuoba Chamb. Claremont.
/'. californica Keys. N. Calif, and Claremont.
P. modica Blackw. Mill Valley, Mt. Shasta.
Sossippus calijornicus Simon. Claremont.
Pirala catifornictis Bks. Mariposa Co.
OxYOPiDAE. Legs long, three tarsal claws, no scopulae. Trochanters not notched.
Eight eyes, dark. Anterior median eyes very small. Abdomen tapers to a joint
behind.
I'eucdia viridans Hent/. Los Angeles.
Oxopes salilcui Hentz. Mill Valley, Palo Alto.
O. rufipes Bks. Mt. Shasta, Santa Clara.
Attidae. Jumping spiders. Short body, Mout legs, two tarsal claws, bright
colors, conspicuous eyes.
Oendryphantes capilatus Hentz. N. Calif.
D. calijornicus Peck. Calif.
D. litis Peck. Claremont.
li. femoratus Peck. Calif.
D. johusoni Peck. S. Calif.. Catalina I., CMareinont.
D. gutlalus Bks. Calif
D. ardfns Peck. Calif.
I), aeneolus Curtis. Palo .Mlo.
[). hartjordi Peck. Claremont.
D. nubitus Hentz. Calif.
Pomona College, Claremont, California 45
D. opifex McCook. N. and Los Angeles Co.
Thiodina retarius Hentz. N. and S. Calif.
falloies signalus Bks. Los Angeles.
P. elegans Peck. San Pedro.
/'. tarsalis Bks. San Pedro.
F. dolosus Peck. Calif.
I', catijornicus Bks. Calif.
P. griseus Peck. Calif.
P. pacifidis Bks. San Francisco.
P. jucundus Peck. Calif.
P. speciosus Bks. Claremont.
/'. hutchensoni Peck. Calif.
Epiblemum palpalls Bks. Palo Alto.
Metacyrba laeniola Hentz. Los Angeles, Claremont.
Marpissa melanognatlia Lucas. N. Calif.
M. catifornica Peck. N. Calif.
Salticus sceniciim Clerk. Santa Barbara L
Atlus dorsatus Bks. S. Calif.
Sidusa morosa Peck N. Calif.
Sitticus claremonti Peck. Claremont.
Sassaciis papenhoei Peck. Calif.
.Utinella dnrsata Bks. Calif.
Pseudiiius siticulosus Peck. Calif.
Hahroifslum morosum Peck. Calif.
Hyctia rohiisia Bks. Calif.
Trap-door spid. ii, 1874, p. 260. Simon List, des osp. 1892, p. 14. Bui. Soc.
Z. Fr. 1884, p. 12, 13, p. 316. Ann. Ent. Soc. Fr. 1883, p. 86, 1891, p. 305, 1893, p. 308.
Proc. Zool. Soc. London. 1880, p. 326, 1883, p. 355. Jour. N. Y. Ent. Soc. 1893, p. 133,
1884, p. 50, 1896, p. 88-110,, 1904, p. 12, 117-118. Ges. Wien. 1871, p. 214. Proc.
Calif. Ac. Sc. 1898, p. 279, 1904, p. 333, 342. Hentz. Spid. U. S. 1875, p. 24, 147.
Verb. Zool. bol. Ges. Wien. 1881, p. 286. Canad. Ent. 1891, p. 209. Cook. Spid.
U. S. 1892. Trans. Conn. Ac. Sc. VI, 1882, p. 9-12, Vlll, 1890, p. 11. Trans. Am.
Ent. Soc. 23, 1896, p. 59-65. Canad. Ent. 1900, p. 97-99, 1898, p. 185. Fuessl. Verz.
D. schw. Ent. Ross. X, 1874, p. 105. Koch Die Arach. VIII, 1849, p 75. Keyserling
Spinn. Am. Thrid. 1884, p. 71—. Proc. Ac. Nat. Sc. Phila. 1878, p. 276, 1888, p. 193,
1892, p. 56, 1901, p. 5-78. Linn. Syst. Nat XI, p. 621. Fab. Ent. Syst. II, 1793, p.
414. Biol. Cent. Am. Arach. 1, p. 51, Spicilog. Zool. 1, 1872, p. 48. Itz. Isis Dresden,
1863, p. 121. Pomona Jour. Ent. VII, No. 3, 1910. Act. Soc. Linn. Bordeaux 1880,
p. 307. Ent. Carnioli 1873, p. 400. An. Soc. Ent. Belg. 1886, p. 56, 1898, p. 25. Bull.
Soc. Zool. Fr. 1895, p. 136. Thorell. Spid. Greenland. 1872. Fab. Ent. Syst. II, 1793,
p. 423. Peckham, Attidae 1883, p. 22. The Entomologist 1894, p. 207. Zoe. 1892, p.
332, 1888, p. 81. Trans. Wis. Ac. Sc. 1900, p. 220. Hist. Nat. d'lles. Canar, 1839, p.
29. Oc. Papers, Wise. N. H. Soc. II, 1895, p. 177. Jour. Ent. Zool. 1915, p. 209, 1916,
p. 112, 1918, p. 1, 1920, p. 1-23, p. 25. Synoptic Index-Catalogue of Spiders of N C.
and S. .'\merica. A. Petrunkevitch Biil. Am. Mus. Nat. Hist. V. 29, 1911.
Ophiuroidea of the West Coast of
North America
ARTHl'R S. CAMPBELL.
This list represenls those Ophiuroidea reported upon by H. L. I lark, J. K. Mc-
Clendon, and others, at various times from the West Coast of North America, ami
especially from the coast of California. Specimens listed are mostly from deeper
water; but a few are littoral.
Original references to each species are given as far as possible. Bathymetrical
ranges given are either extremes or are the only point from which specimens are
known.
There seem to be several restricted faunas represented in the list. It is quite
possible that specimens of almost any of the list might he taken at other points off
the coast, and thus extend, the known range.
The purpose of the list is to clear up certain synonyms, to check the present
literature so far as possible, to record more complete data concerning the distribution
of forms likely to be taken nearby, and to know more thoroughly what we have.
Our work is by no means finished, but we feel the list may he of some aid to
those undertaking the study of west coast forms.
Ophiurae
Ophiodermatidae.
Opiiioderma panamensis Liitkin. Add. ad Hisi. Oph., 1, p. 193. 1859. Littoral.
Panama to California.
Ophiodcrma furift/nta Liitkin. 1S59. Add. ad Hisi. t)ph., 2, p. 21. Littoral.
Lower Calif.
Op/iiocryptus maculosus Clark. 1915. 3d. Laguna Rep., Pomona Coll., p. 64.
Littoral. Laguna, Calif.
DiopeJfrma axiologum Clark. 1915. Ech. Lower Calif., p. 206. pi. XLV, fig.
5-7. Am. Mus. N. Hist., vol. 22, art. S, pp. 185-236. Coast. Cape St. Lucas.
Ophiolepidae.
Ophioplocus tsmarki Lyman. Bull. \\. C. Z. 3, pt. 10, p. 227, p. 5. Shore-4il
faths. Panama — north.
Ophiocten pacificum 1.. k M. Mem. M. C. Z., 23, no. 2, 18S7. 0-1573 faths. San
Diego southward.
Ophiomusium jollirnsis McClendon. V . C. pub. Zoo., vol. 6, no. 3, p. 36. 1909.
La Jolla, Calif. 85-330 faths.
Ophinmusium lymani \V. Thos. "Pep. of the Sea", p. 172, figs. 32-33. 600-1,101
faths. Cosmopolitan.
Ophiomusium ylahrum I.. & M. Mem. M. C. Z., vol. 23, p. 132. 480-2,232 faths.
Kquator-47° N.
Ophionrrris adsprrsus Lvman. Bull. M. C. Z., vol. 10, p. 236. 647 faths. Lower
Calif.
Ophionrreis pnlypnrus L. k M. Mem. M. C. Z., vol. 23, p. 109. 491-647 faths.
Lower Calif.
Pomona College. Claremont, California 47
Ophionereis annulala Le Conte. Proc. Acd. N. Sc. Phila., p. 317. 1851. Shore-
35 faths. California.
Ophiura flagellaln (Lyman) Meissner. 1901. Das Thierreich, vol. 2, pt. 3, p.
925. "35 faths. Lower Calif.
Op/iiura superba (L. & M.) Meissner. 1901. Uas Thierreich, vol. 2. pt. 3, p. 925.
Lower Calif.-northward. 451-930 faths.
Ophiura irrorata (Lyman) Meissner. Das Thierreich. vol. 2, pt. 3, p. 925. 1,760
fath^. Lower Calif.
ophiura ponderosa (Lyman) Meissner. 1901. Das Thierreich. vol. 2, pt. 3, p.
925. 640 faths. Lower Calif.
Ophiura ogliopora Clark. Ech. Lower Cal., p. 210, pi. 45, figs. S-9. M. N. Hist.,
pp. 185-236, 1913. 630 faths. Cape St. Lucas.
Ophiura sarsii Liitkin. Vid. Medd. for 1854, 1885, p. 101. 5-695 faths. Cosra.
Ophiura leploclenia Clark. Bull. U. S. N. M., no. 75, p. 51. 1911. 67-1,771
faths. Northward.
Ophiura cryptolepis Clark. Bull. V. S. N. M., no. 75. p. 69. 1911. 230-636
faths. Northward.
Ophiura lulkiui Lyman. Proc. Bost. Soc. N. Hist., 8, p. 197. 1860. California to
Puget Sound. 22-600 faths.
Ophiura kofoidi McClendon. U. C. pub. Zoo., vol. 6, no. 3, p. 38. 1909. SO
faths. San Diego.
Ophiura hrevispina (Say) Lyman. "Challenger", Zoology, vol. 5, p. 9. Deep
Water. Puget Sound.
Amphiuridae.
Amphiodia barharac Lyman. 111. Cat. M. C Z. Harvard, 8, pt. 2, p. 17, pi. 3.
Shore-lOO faths. Deep in sand. California.
Amphiodia sirnni/yloplax Clark. Smith. Bull. 75. 1911. p. 164. 171 faths.
Washington.
Amphiodia urliai Lyman. Proc. Bost. Soc. N. Hist., vol. 7, 1860. p. 195. 15-50
faths. Calif-Alaska.
Amphiodia occidentalis Lyman. Proc. Bost. Soc. N. Hist., vol. 7, 1860. p. 194.
Coast. Monterey-Alaska.
Amphiodia periercta Clark. Smith. Bull. no. 75. 1911. p. 160. Oregon-Alaska.
8-240 faths.
Amphiodia daira Lyman. 1879. Bull. M. C. Z., vol. 6, p. 27. 1,076-1,760 faths.
North.
Amphiodia curyaspis Clark. Bull. V. S. N. M. no. 75, p. 158. 68-318 faths.
North.
Amphiura diastra McClendon. U. C. pub. Zoo., vol. 6, no. 3, supp. San Diego.
100 faths.
Amphiura tarihara Clark. 1911. Bull. 75, U. S. N. M., p. 142. 1,090 faths.
Northward.
Amphiura diomedeae L. & M. 1899. Mem. M. C. Z., vol. 23, p. 151. 640-659
faths. Monterey-Southward.
Amphiura serpentina L. & M. Mem. M. C. Z., vol. 6, p. 143. 1899. 475-645
faths. North.
48 Journal of Entomology and Zoology
Amphilimna pentacantha Clark. Smith. Bull. 75. 1911. p. 172. 4S faihs.
Calif.
Amphipholis pugtiana Lyman. Proc. Host. Soc. N. Hist., vol. 7. 1868. p. 193.
8-240 faths. Monterey-North.
Amphipholis punlarenae Liitkin. Bidrag til Kundskab. cm Slagestjerne, 3 Vidensk.
Meddel. Naturhist. Foren : Kojobenh. 1856. La Jolla. 10-50 faths.
Ophiocnida hispida Le Come. Proc. Acad. N. Sc. Phila., 5, p. 318. 1S51. Shore.
Panama-Catalina.
Ophiocnida amphacantha McClendoii. V. C. pub. Zoo., vol. 6, no. 3. 1909. p.
46. 120-150 faths. San Diego.
Ophiopholis aculeata Linn. Syst. Naturae, 12th Ed., 1767, p. 1101. 9-372 faths.
Puget Sound-North.
Ophiopholis aculeata kennerlyi Lyman. Proc. Bost. Soc. N. Hist., vol. 7, 1860.
p. 200. 8-238 faths. Calif.-Alaska.
Ophiopholis bakeri McClendnn. l". C. pub. Zoo., vol. 6, no, 3, p. 41. Southern
Calif. 60-215 faths.
Ophiactus arenosa Liitkin. Bidrag til Kundskab om Slagestjerne, 3, Vidensk.
Meddel. Naturhist. Foren: Kojobenh. 1856. in sponges, Lower Calif. -South.
Ophiocomidae.
Ophiocoma arlhinps Liitkin. 1859. Add. ad Hist. Ophiu., pt. 2, p. 145. Coast.
Lower Calif.
Opiocoma aUxandri Lyman. Proc. Bost. Soc. N. Hist., vol. 7, p. 256. Coast.
Lower Calif.
Ophiopleris papillosa Lyman. 111. Cat. M. C. Z., 8, pt. 2, p. 11, 1875. Shore-3ii
faths. California.
Ophiocanthidae.
Ophioianiha rhathnphora f'lark. Smith. Bull. no. 75, p. 201. 451-630 faths. Ber-
ing Sea-Lower Calif.
Opiocaniha normani Lyman. Bull. M. C. Z., 6, no. 2, p. 58. 1851. 600 faths.
East and West Pacific.
Opiocaniha hairdi Lyman. 1883. Bull. M. C. Z., vol. 10, p. 256. 451-525 faths.
North.
Opiocaniha halhyhia Clark. Bull. I". S. M. no. 75, p. 232. 1911. 868-1,090 faths.
West Pacific.
Opiocaniha monilijnimu 1.. Si M. 1899. Mem. M. C. Z., vol. 23, p. 171. 284
faths. Panama-Lower Calif.
OPHIOTHRICIDAE.
Ophiolhrix spiculala Lc Conle. Proc. .'\cad. N. Sc. Phila., v.. p. 318. 1851. Shore
— lOOfaihs. Alaska-Panama.
Ophiolhrix nidis Lvman. Bull. M. C. Z., pt. 10. p. 239. 1874. Shore— La Jnlla.
OPHIOMV-VIDAE.
Ophiocynodus coryncles Clark. Smith. Bull. 75. p. 274. 345-685 faih>. Wavh-
ingtnn.
Pomona College, Claremont, California 49
EURYALAE.
ASTEROCHEMIDAE.
.Istrochema sublaeve L. & M. 1899. Mem. M. V. Z., vol. 23, p. 187. 534 faths.
Lower Calif.
.Uleronyx dispar L. & M. 1899. Mem. M. C. Z., vol. 23, p. 185. 491-1101 faths.
Lower Calif.
.■\STER0PHVT1DAE.
Asleronyx excavala L. & M. Mem. M. C. Z., vol. 23, p. 185. 491-525 faths.
lower Calif.
hleronyx Itiveni M. & T. 1842. Syst. .Ast., p. 119. 284-659 faths. North-Lower
Calif.
Cnrgoncephalus eucnemis M. & T. Syst. Aet., 1842. 160 faths. Laguna-North.
Gorgoncephalus caryi Lyman. Proc. Bost. Soc. N. Hist., vol. 7, 1860, p. 424. 8-576
faths. San Francisco-Northward.
(Contribution from the Xoolo'/ical Laboratory of Pomona College.)
VII. Round Worms
Nematoidea. The central nervous system of nematode worms
was early described as a whole by Biitschli who recognized a collar
of nerve cells and fibers and longitudinal strands. Hesse, 1892,
gives a clearer picture of the nervous system of Asccdis and others
since that time have improved and elaborated upon these and other
early suggestions. Especially noteworthy are the works of Gold-
.schmidt, 1908-9, and Deineka, 1908, each very valuable although
the two investigators disagree on many points.
The nervous system of Ascaris may furnish a good starting
point in a discussion of the nervous system of the group. In this
genus there is a circumoral ring about the pharynx near the anter-
ior end of the body. Ganglion cells are not abundant. They are
chiefly grouped about the origin of the nerves. The nerve ring gives
off six or more longitudinal nerves of which the mid-dorsal and mid-
ventral are usually the largest and are connected to each other by
fine branches. At the caudal end the lateral nerves pass into two
branches formed by the division of the ventral nerve. Just above
this point the ventral nerve swells into the anal ganglion. In the
male the anal ganglion gives oflT two lateral nerves which form a
ring about the cloaca.
The nerve ring forms a plexus according to Goldschmidt, in
that all fibers are connected to other parts, but the plexus is regular
and not of the diffuse type as found in Coelenterata. Three cell
types are found, sensory, association and motor. Besides the direct
connection of cell with cell through their processes there is in places
a true neuropile. Neuroglia cells are found but are not prominent.
Deineka favors the neropile method of interrelation more than Gold-
schmidt. This author also has demonsti'ated the neurofibrillar
arrangement of the material with the nerve cells and has shown
rather elaborate interrelations between the fibrils of associated
cells. He shows nerve terminations in muscle and sensory endings
in the skin of the body. Aside from the general surface of the body
the three papillae about the mouth are the only sense organs. These
are supplied by six short nerves running from the nerve ring.
With free living nematodes but little has been done. In Enoplas
Hilton, 1920, a very marked head ganglion above the mouth has two
strands running backwards to the thick mid-ventral nerve strand
and from the dorsal side a slender dorsal nerve runs the length of
the body. The ganglion is rather complex in structure. From an
inner group of nerve cells, fibers run forward to the sensory epithe-
lium of the tip of the snout and three eyes, one dorsal and two
ventro-lateral are composed of pigment and clear area in front.
Magrath, 1919, in Callanus, gives a good account of the nervous
system of this simple nematode.- In this as in other forms, there is
56
NERVOUS SYSTEMS AND SENSE ()R(JANS
FiR. 15. A. Diagram of the nervous system of Axcari.s, after Hesse. B. Dia-
Rram of the nervous system spread out flat, from Goldschmidt. C.
Plan of the central nervous system of Ancarix, after Deineka. D H.
Sensory terminations and peripheral nerves of Asrarin, after Deineka.
ROUND WORMS
57
a cephalic commissure. With this are associated twenty nerve cells
on each lateral half and a large number just anterior to it. From
these last groups six slender nerves pass forward close to the oeso-
phagus to supply the anterior region. The two sub-ventral have
small ganglia upon them. Connected with the caudal edge of the
nerve ring are four chief ganglia, one dorsal, one ventral and two
lateral. Each of these has long strands extending tow-ards the tail
Fig. 16. The figure above is a reconsti'uction of the head end of Enophis,
showing the position of the nei'vous system. The lower figure at the
left is of a section through the whole body of the worm, showing the
dorsal and ventral nerve bands. Both these figures enlarged 75
times. The drawing at the right is from a section through the head
ganglion, enlarged 170 times. The dorsal side is up in all the figures.
Hilton.
end of the animal. Continued from the ventral and separated a
little distance is another ventral ganglion, the post- ventral. The
dorsal cephalic ganglion is the smallest; the lateral cephalic ganglia
are the largest. As pointed out by others the cephalic commissure
or nerve ring is essentially fibrous. The fibers are derived from the
ganglia connected with it.
58
NKKXOUS SYSTEMS AM) SENSK ORCJANS
In the female the central anal ganglion is the largest. It con-
nects with smaller lumbar ganglia out laterally and by a loop with
the recta! ganylion.
In the male the anal ganglion is large, but the two lumbar are
nearly as large. Two rings of nerves are connected with the anal
ganglion and one with the small cloacal, and the other with the
rectal ganglion.
GORDiODEA. Villott, 1874, shows that the ventral cord repre-
sents the central nervous system with an anterior and po.sterior
V
?
Fife. 1". A-F. Nervous system of Goidoidea. A. Section through brain and
subocsophepeal band, much chanped from Montgomery. B. Petition
of supra and suboesophapeal panplion modified from Montpomery.
C. and D. Sections of vintr-il cord. E. and F. eras'; and longitudinal
sections of the ventral cord after May. G and H. Head of Chaetog
iinlhn after Hertwip, showinp brain, sense orpans and chief nerves.
I. Ventral panplion shown, Hertwip. J. Eye of Chactognatha. K.
Ganglion in body of Aciii'li'ircphalia after Leuchart.
ganglion. In 1887 he traced fibers from the head ganglion into the
thickened hyrodermis of the head. Vejdovsky, 188,3, 1894, con-
siders that there is no cerebral ganglion and no ganglion cells on
the dorsal side of the peripharyngeal ganglion. He distinguishes
neuroglia cells.
ROUND WORMS 59
Ward, 1892, on Nectonema, a pelagic marine form, gives an
account of the nervous system. The anterior ganglionic mass or
brain forms a large portion of the floor of the anterior chamber.
The oesophagus lies in a groove in its center. There is but a slight
dorsal commissure above the oesophagus. The ganglion cells are
not abundant in the brain. A smaller kind is more abundant than
another sort which is very much larger. There are five pairs of
these last which are nearly constant in position and form. The
ventral nerve cord continues from the brain and runs the length of
the body separated into three ai'eas to correspond to the three nerves
of which it is composed. Some large cells in the cord are much like
those in the brain. In the male the ventral cord is much enlarged,
being larger than the brain itself. In the female the anal ganglion
is but slightly larger than the central cord with which it is
connected.
Camei'ano, 1897, considers the nervous system to consist of a
supraoesophegeal ganglion and a ventral nerve strand. Mont-
gomery, 1903, finds a ventral unpaired nerve trunk with the cephalic
ganglion at its anterior enlargement and the caudal or cloacal
ganglion, a posterior enlargement. To the peripheral nervous
system belong the neural lamella; the endings in the hypodermis of
the fibers of nerve cells situated in the central nervous system ; the
hypodermal longitudinal nerve ; sensory cells in hypodermis ; non-
sensory hypodermal nerve cells and the nerve fibers which innervate
the cloaca of the female and the vasa defFerentia of the male. Two
types of cells were found in the nerve cord. One type contained but
little chromatin. These cells on the lateral sides of the cord are
quite uniform and small. On the ventral side there are smaller and
larger cells of this type. The larger or giant cells are less numer-
ous. Sometimes there is a paired arrangement of these cells but
usually they are irregularly placed one behind another. These
cells seem to be bipolar with two large processes proceeding from
the cell directed towards the fibrous core of the nerve cord. Some
of the small cells appear to be bipolar or multipolar. All cells are
without membranes. Montgomery thinks that these deeply staining
cells are probably motor and visceral in function.
The deeoly staining cells seem to be multipolar with very long
nrocesses. It could not be determined whether there was anas-
tomosis of the processes. These cells seem like the multipolar
neuroglia cells of other invertebrates but processes pass into the
hypodermis.
The ventral cord seems to be made up of three converging rays
of fibers but each lateral ray is made up of several distinct fiber
tracts. The median tract is the largest and is made up of longi-
tudinal fibers which are closely arranged. Very rarely are nerve
60 NERVOUS SYSTEMS AND SENSE ORGANS
cells found on the dorsal side of this tract. They are most abundant
at its ventro-lateral angles.
On each side of the median tract are three not sharply marked
portions; (a) a dorsal tract mostly of deep staining fibers, (b) a
latero-ventral tract bounded by a layer of clear cells, a tract
mainly made up of longitudinal dark fibers, (c) a medio-ventral
tract larger than the last and between it and the median. It con-
tains dark fibers running in all directions but mainly longitudinally
and also clear fibers.
The nerve cells send their fibers in radially. The "Punktsub-
stanz" is composed of fibers from two kinds of nerve cells.
The nerve cord has no neural sheath but is immediately sur-
rounded by a small-celled parenchyma. Outside of the outer nerve
cells of the cord is a sheet of dark staining fibers.
At intervals along the nerve cord are transverse commissures
of fibers e.xtending from the dorso-lateral angle of one side to that
of the other. There is no segmental grouping of the nerve cells.
The transverse commissures also are not nietameric as they are too
irregular and too close together.
The so-called cephalic ganglion is a slightly enlarged anterior
end of the nerve cord. It is more thickened from side to side than
dorso-ventrally. The nerve cells are numerous but limited to the
median line. In the head the fiber tracts appear like a large median
one each side of the middle line. There is a transverse commis-
sure near where the cephalic nerves meet. As this is on the ventral
side it has been called the ventral commissure. According to Mont-
gomery there is no brain or supra-oesophageal ganglion.
The cloacal ganglion of the female is the enlarged posterior
end of the ventral nerve cord .just anterior to the point where the
lateral lobes branch. From the ganglion there are anterior and
posterior cloacal nerves.
The cloacal ganglion in the males is not so sharply limited as in
the female. The length of the ganglion varies in different indi-
viduals of the -same size. Small nerves pass to the vasa deferentia.
The ganglion divides into a right and left caudal nerve into the
caudal lobes.
In both sexes the neural lamella attach the nerve cord to the
hypodermis. It is it.self of hypodermal nature. At the point of
the attachment of the neural lamella, the hypodermis is conical on
cross .section. There is a clear area here in which the longitudinal
hypodermal nerve is located. It is comiio.sed of nerve fibers from
dark nerve cells of the ventral cord. This hypodermal nerve runs
as far as the central nervous system.
Fibers enter the hypodermis by way of the neural lamella
apparently from cells in a ventral position. Upon entering the
ROUND WORMS 61
hypodermis some run longitudinally in the hypodermal nerve or
along the sides of the body.
There are two main types of sensory cells in the hypodermis,
small irregular cells staining deeply and the elongated cuticular
cells of the mid-ventral line. Motor cells are considered to be the
clearer ones of the nervous system, the darker straining cells the
sensory ones. These last run into the hypodermis.
Linstrow, 1889; Ward, 1892; and Montgomery, 1897, have
found structures in the anterior part of the head which may be an
eye or possibly a part of the head ganglion.
May, 1919, recognizes more clearly than Montgomery a ring
of nervous tissue in the head region. In Gordius the brain is out-
lined at the first as a ring of cells in the hypoderm of the proboscis.
It soon separates remaining connected only at the anterior end and
ventral side. At first it consists of a few large cells which surround
the larval muscles. These large cells remain in this position while
the rest of the brain develops in front. The ventral cord arises as
a thickening of the hypoderm, but later separates from it. The cells
that make up the nerve cord at first appear as two rows of nuclei
on the ventral side of the larva. The larger cells seem to be bipolar,
giving ofl: one fiber to the longitudinal tract and one to the dorsal
border of the cord.
The brain of Paragordius develops later than that of Gordius.
In the first genus the cells of the lamella are located in the ventral
cord while in Gordius it consists of a series of cells. According to
May the mass of cells which Montgomery calls retina is the larger
pai't of the cephalic ganglion.
The reactions of gordioid worms is slow and of a primitive
nature. The grasping reaction of the male when in contact with the
female is the most definite. If a specimen is at rest it usually re-
quires several successive stimuli to cause even a slight movement of
the body. There seems to be no distinct response to light.
ACANTHOCEPHALIA. In this group the nervous system is found
to be a single ganglion of large cells located on the surface of the
proboscis near its base and two small ganglia in the male which
supply the reproductive organs. The larger cephalic center gives
off nerves to the proboscis in a cephalic direction and through the
lateral retractor muscles on each side caudally strands run out to
supply the body-wall. There are no sense organs known.
Chaetognatha. In Sagitta the nervous system consists of a
cerebral ganglion in which eyes ai'e situated. A large ventral gang-
lion is situated about one-third or one-half of the way down the
body. Oesophageal conectives join these two chief ganglia. Fibers
run from the head ganglion to the jaws and sense organs of the head
region and two other small ganglia have been described near the
62
NERVOUS SYSTEMS AND SENSE ORGANS
mouth. From the large ventral ganglion many branches run to
lateral and caudal regions of the body. This ventral ganglion is the
chief one from the standpoint of size.
Many papillae on the surface of the body probably serve as
organs of touch. The eyes, one on each side of the dor.sal region
of the head are globular and each contains three biconvex lenses
separated by pigment and surrounded by rod-like sensory cells.
About the dorsal part of the head end there is a ring-like ridge
bearing modified ciliated cells. This has been called the olfactory
ring.
In Sagitta, a great proliferation of cells in the head region of
P"ig. 18. The sketch at the right is an outline of a larval Sagitta showing
the position of the origin of the two chief ganglia and the lateral
sense organs. All are indicated by the darker shaded areas. The
figure at the left shows the position of the chief head ganglia of
Sagitta.
the elongated larva forms the brain. This is added to on each side
by two lateral ridges which later unite to form the cephalic hood.
The ventral ganglion begins as a thickening t)f the ectoderm from
behind the head alx)ut two-thirds of the length of the body. A tac-
tile organ is developed from ectoderm on each side of the tail region
ROUND WORMS 63
a little distance from its end. At a latei' time a double curved line
of nuclei forms a horse-shoe shaped area, the so-called olfactory
organ.
BIBLIOGRAPHY
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1874. Beitrage zur Kenntnis des Nerv(;ns\ stems der Kematoden. Arch.
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64 NERVOUS SYSTEMS AND SENSE ORGANS
Krohn, A.
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ROUND WORMS 65
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135-188, pi. 8.
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JOURNAL OF
ENTOMOLOGY AND
ZOOLOGY
VOLUME XIV. 1922
PUBLISHED QUARTERLY BY THE
DEPARTMENT OF ZOOLOGY OF POMONA COLLEGE
CLAREMONT. CALIFORNLA, U. S. A.
CONTENTS OF VOLUME XIV
Volume XIV. Number 1
Alexander, Charles P.
The Biology of the North Aimri-
can Crane-Flies. 1.
Chamberlin, Ralph V.
A \\\v I'laty(lrsiii()i<l Diplupod
friiin California, H.
Gampbell, Arthur S.
Hydroids Niar LnKUna Beach. Id.
Hilton, W. A.
Xcrvous Svstcni and Sense Or-
gans. VHl. ]S
Volume XIV, Number 2
Penny, Donald D.
A Catalog of the California Alcy-
rodidae and the Descriptions of
Four New Species. 21.
Campbell, Arthur S.
rnliniinary N'otcs on (_iro\vtli-
.Stagcs in lUittk-Stars. 37.
Hilton, W. A.
Xcrvous System and Sense Or-
gans. ^5.
Volume XIV. Number 3
Marimon, Sarah
The Skull ol .N'otothalanius Toro
sus. 55.
Essig, E. O.
.\ N'cw .Xphis on California Sam
Hilton, W. A.
I'horonida and .\ctinotrocha. 65.
Volume XIV. Number 4
Hilton, W. A.
Tlu- Occurnnn' nl l'olynordin>
.•\dnlt at Lagnna Beach, 7i.
Essig. E. O.
Insect Xotis from Laguna lUacli.
California. 75.
Hilton, W. A.
Xcrvous Svstcni and Sense Or-
gans. XI.' 79.
INDEX TO VOLUME XIV
Actlnotroclia, 65. Hydroids, 10.
Alexander. C. P.. 1. Hymcnoptera. 78.
Aleyrodidae, 21. Insect.s, 7.5.
Bracliiopoda. 79. l.epidoptera, 77.
Uryozoa, 45. Marimon, S., 55.
Campbell, .\. S., 10. ^7. Ncrvou.s System. IS. 45. 65. 79.
Chaniberlin. R. V.. 8. Notothalamu.<;. 55.
(rane-Flie.s. 1. Orthoptera, 75.
Oiplopod. 8. reniiy. D. D.. 21.
Diplera. 77. I'lioronis. 65.
Essig, K. (_).. 61. 7,1 I'ldyKordius, 7.?.
Hemiptera, 76. .Sen.se Organs. 15. 45, 65, 79.
Hilton, W. A.. 15. 45. 65. 73, 79. Serpent Stars, 37.
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JOURNAL
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MARCH, 1922
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT o/^ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
The Biology of the North American Crane-Flies
Charles P. Alexander --..--.-- 1
A New Platydesmoid Diplopod from California,
Ralph V. ChamberUn - - 8
Hydroids Near Lacuna Beach, Arthur S. Campbell - - 10
Nervous System and Sense Organs, VIII, W. A. Hilton - 15
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The Biology of the North American
Crane-Fhes
(Tipulidae, Diptera)
VI. The Genus Cladiira Osten Sacken
By Charles P. Alexander
Generic Diagnosis
Larva. Form comparatively short and stout ; integument
provided with a delicate appressed pubescence: no distinct setae;
basal annulus of each of abdominal segments two to seven with a
transverse area of microscopic points arranged in long, transverse
rows. Last ventral segment with a Hattened lobe covered with
short setae, evidently an organ for shoving. Spiracular disk
entirely without lobes, the spiracles being situated on the exposed
dorso-caudal surface of the last abdominal segment. Head-capsule
relatively compact : frontal plate broad, only slightly narrowed
behind. Labrum quadrate, with conspicuous, oval, lateral arms ;
antennae two-segmented, the terminal segment elongate-oval :
mandibles of a herbivorous type, with an apical point and two
incomplete rows of teeth on the inner or cutting face; mental bars
widely separated, each bar provided with two acute teeth at its
mesal end.
Pupa. Cephalic crest gibbous, entire or feebly bifid, armed
on either side with a single powerful bristle ; two bristles on both
the front and vertex: labrum with pair of small bristles at each
cephalic-lateral angle ; labial lobes subquadrate, weakly separated
by the apex of the labral sheath : palpal sheaths short and stout,
straight ; lateral margins of eye produced laterad into a digitiform
lobe : antennal sheaths extending to opposite one-third the wing-
sheaths. Pronotal breathing horns lacking, entirely sessile ; pro-
notum and mesonotum armed with conspicuous bristles : wing-
sheaths ending opposite the base of the third abdominal segment ;
leg-sheaths long, ending opposite the base of the sixth abdominal
segment, the hind legs longest, the middle legs shortest. Abdominal
tergites with ten strong bristles, eight being arranged in a single
transverse row along the posterior margin ; abdominal pleurites
with four strong bristles, one on anterior ring, two near the caudal
margin of the posterior ring, one ventrad of the spiracle ; spiracles
rudimentary, situated on segments two to seven ; sternites unarmed
with bristles.
Discussion of the Genus
The genus Cladura was erected by Osten Sacken in 1859
(Proc. Acad. Nat. Sci. Phila., p. 229). The genus includes but six
2 Jiiurn.il of Entomolotiy and Zoology
known species, with a Holarctic distribution, there l)eing two
species from eastern North America, one from welter i
North America, and three from Japan. Of the eastern North
American species, the most common and best-known is the geno-
type, Cladura flaroferruginca. The six known species of the genus
are all forms that appear on the wing in late summer and in
autumn.
The only reference to the immature stages of this curious
genus is the brief diagnosis by the writer (The Crane-flies of New
York, Part II. Biology and Phylogeny. Cornell University Agri-
cultural Experiment Station. Memoi'r 38. p. 9)9 ; lf)2i ). The geno-
type is common and widely distributed throughout the northeastern
United States, but until the ])resent year the writer had been
unable to locate the immature stages. The conditions under which
these stages occur are briefly outlined herein.
Augurville, or Brownsfield. Woods, near Urbana. Illinois, is
an open, low Transitional or upper Austral woodland, traversed
in spring and early summer by a small stream. In early
spring the valley through which this brook flows is car-
peted with a dense growth of Blue-eyed Mary {Collinsia vertia).
On the higher ground and dry slopes, other characteristic
spring flowers, such as TrilUtn)i recnrratum. Claytonia
virginica. squirrel-corn, dutchman's breeches, blood-root, white
trout-lily, and other forms, occur in numbers. The forest cover
consists of linden, hard maple, buckeye, hackberry. bur oak, honey-
locust, and a few less common species, certain individuals of all of
these species being giants of their kind and evidently members of
the primitive forest. The undergrowth consists principally of
pawpaw and s]iice-bush, together with considerable rejiroduction
of buckeyes and other trees. In the autumn, the vernal flora is
replaced by the dominant wood-nettle, many species of Aster and
Snlidaf/o. some Eupatorhim and other late summer plants. Adults
of Cladura flnrnfcrruginea were found in these woods during the
fall of 1919.
On September 5, 1920, Mrs. Alexander and the writer began
a systematic search for the larvae of Cladura. Earlier experience
in Maine, New York, and Kansas had demonstrated that it was
highly improbable that the early stages were to be found in mud,
or even in damp earth, or in decaying wood, these habitats being
those commonly frecjuented by the early stages of the Tipulidae. A
careful search was instituted in soil that was liaked comiiarativoly
hard and dry. The lumps were dug out and crumbled into dust, the
contents being carefully examined. This method of search soon
revealed a short, stout, light yellow crane-fly larva, that was at
once determined as probably l)eing that of Cladura. On this date,
the only other insects associated with this larva were larvae of the
Scarabaeid, Xnlorijctis .■<atitru.-< (Fal)r.), a Tenebrionid, Miracau-
tha contrarta (Beauv.). and a few adult Corabidae and Stajihy-
Pomona College, Claremoiit. California 3
linidae. The conspicuous millepede Spirobolus marginatum
(Say), was also found in these situations. The soil was covered
with a layer of dead leaves and other vegetable detritus, but this
had not been sufficient to prevent the dessication of the soil to a
depth varying from six to twelve inches or more. Three larvae
taken on September 5 were placed in breeding vials.
On September 19, 1920, Mrs. Alexander and the writer con-
tinued the search in these same haunts, and this resulted in the
discovery of six additional larvae and four teneral pupae. As
before, they occurred in soil that was very dry, underneath a layer
of leaf-mold and other debris. These were placed in rearing.
On September 29, 1920 the writer again went to Augurville
Woods. The weather was very cold and raw. By careful search-
ing, eight pupae were discovered, some being very dark colored and
evidently nearly ready to transform to the adult condition. These
were placed in tin salve boxes for rearing. On the following day,
two females of Cladura flavoferruginea emerged from two of the
pupae discussed above. Other adults emerged during the following
week. The remaining larvae and pupae were preserved in alcohol.
Bergroth and other writers had surmised the relationship of
Cladura to the nearly apterous snow-fly, Chionea Dalman, a fact
that is amply substantiated by the discovery of the larvae of the two
genera. Brauer, Egger and Frauenfeld (1854) had taken gravid
females of the commonest European species of Chionea, C. aran-
eoides. and confined them in breeding jars, where they laid a
large number of eggs, which hatched into stout yellow larvae that
agree in many features of their organization with the larvae of
Cladura described in this paper. Unfortunately the larvae of
Chionea have never been carried through to the pupal condition.
The larvae of the two genera agree in their short, stout form,
the obliquely truncated spiracular disk that is quite devoid of sur-
rounding lobes, and in the general features of the head capsule. The
pupa of Cladura is notable by the entire lack of protuberant breath-
ing-horns, the breathing-pores being entirely sessile. The nearest
approach to this condition in the Tipulidae is found in the genus
Dieranoptycha Osten Sacken, which is likewise characteristic of
unusually dry conditions in open upland woods. Other notable
features of the pupa of Cladura are found in the very elongate leg-
sheaths and the unusual development of long setae on the dorsal
and pleural regions of the abdomen. The pupa is very small com-
pared with the adult which emerges from it.
Natural Affinities
The genus Cladura unquestionably belongs to the tribe Eriop-
terini where it was placed by Osten Sacken. The discovery of the
immature stages confirms the belief that this genus, as well as
Chionea Dalman, and probably Crypteria Bergroth and Ptero-
4 Journal of Knt()m(il<)j;\ and Zoolojiy
chionea Alexander, should be isolated from the Eriopteraria where
now placed and made a separate subtribe, the Chionearia or
Claduraria, the former name being based on the oldest genus.
DESCRIPTIOxN OF THE IMMATURE STAGES
Larva — Length (fully grown), 10-10.5 mm.
Diameter, 1.2 mm.
General coloration light yellow throughout.
Form comparatively short and stout. Integument provided
with a delicate appressed pubescence; no di.stinct setae. Abdomi-
nal segments divided into a narrow basal annulus and a much
broader posterior annulus, the latter being approximately two and
one-half times as long as the former ; the ventral surface of the
basal annuli of abdominal segments two to seven with a conspicu-
ous transverse area of microscopic foints arranged in long trans-
verse rows.
Spiracular disk entirely destitute of lobes, the spiracles being
located on the obliquely truncated dorso-caudal surface of the last
abdominal segment. Spiracles circular, the ring pale, the centers
dark; spiracles separated from one another by a distance about
equal to or a little less than the diameter of one. Ventral surface
of the terminal abdominal segment with a projecting, flattened
lobe that is provided with a dense brush of short, pale setae, th's
organ presumably being used for propelling the insect through
the soil.
Head entirely retractile. Head-capsule very compact for a
member of the Eriopterini. Frontal plate broad, only slightly nar-
rowed behind, the apex obtuse or subtruncate. Labruni-epiphar-
ynx quadrate, the surface covered with short, dense hairs; on
either side a stout ova! arm or lobe directed cephalad, these arms
connected by narrow bars, with the frame-work of the head. Men-
tal bars entirely separate, each bar with two acute teeth on the
cephalic side immediately before the apex. Antennae two-seg-
mented, the basal segment short-cylindrical, the terminal segment
elongate-oval, gradually narrowed to the obtuse ajiex. Mandibles
relatively s'eiuler. of a herbivorus tyi e, tlie teeth blunt: anical
point small ; two incomplete rows of flattened obtuse denticles
along the inner face of the mandible, the outermost tooth of each
row largest, the others gradually smaller, becoming subobsolescent ;
the basal teeth are very tiny, arranged in short combs; proximal
caudal angle of the mandible produced into a cylindrical chitini/.ed
])ar. Maxillae consisting of simple hairy lobes.
Pi(j)(i — Length, 6.7 mm.
Width, d.-s., 1.4 mm.
Depth, d.-v., 1.4 mm.
Pomona College, Clnrenioiit, California 5
The coloration of newly transformed papae is pale yellow. In
older individuals, the thorax, head and sheaths of the appendages
gradually deepen in intensity to almost black in specimens about
to transform.
Cephalic crest projecting between the antennal bases as a
gibbous lobe that is entire or microscopically bifid, on either side
with a conspicuous erect bristle situated immediately dorsad of
the base of the antenna. Vertex between the cephalic ends of
the eyes with a strong bristle on either side, immediately caudad
of each of which is a small tubercle. Frontal region likewise with
a pair of strong bristles that are somewhat appressed against the
face, directed caudad. Labral sheath with the apex rounded, very
narrowly separating the labial lobes ; at the base of the labrum iv^
either side are two small bristles ; sheaths of the palpi short but
stout. Lateral margin of the eyes produced laterad and slightly
caudad and dorsad into a conspicuous finger-like lobe. Antennal
sheaths extending to about opposite one-third the length of the
wing-sheath.
Pronotal breathing horns entirely lacking, the pores being
sessile, lying immediately dorsad of the antennal sheaths. Prono-
tal scutum with two weak bristles behind the antennal sheaths ; pro-
notal scutellum with three powerful bristles on either side near the
summit. Mesonotum gibbous but unarmed with tubercles or spines.
The following mesonotal bristles are evident : one on the ventral
caudal angle immediately cephalad of the wing-root ; a group of two,
one being much smaller than the other, immediately at the wing-
root ; a transverse row of three strong bristles on either side, dor-
sad and proximad of the wing-root ; two weak bristles slightly ceph-
alad of the level of these latter three, one on either side of the me-
dian line; a strong bristle dorsad and cephalad of the pair at the
wing-root. Metanotum with a strong bristle at the ventral cephalic
angle. Wing-sheaths extending to opposite the base of the third ab-
dominal segment. Leg-sheaths long, extending to opposite the base
of the sixth abdominal segment ; sheaths of the posterior legs long-
est, a little exceeding those of the fore-legs ; middle legs shortest,
ending immediately beyond the base of the last segment of the
posterior sheaths.
Abdominal tergites and pleurites with very conspicuous bris-
tles ; sternites entirely unarmed. The distribution of the setae is
as follows : On the tergites — no setae on the anterior annulus ; on
the posterior annulus a single transverse row of eight long bristles
along the posterior margin of the segments, four on either side of
the median line ; cephalad of the outermost pair of these bristles
and located on the anterior part of the posterior annulus is a single
strong bristle on either side: on the eighth tergite there are only
four bristles, arranged to form a rectangular or trapezoidal figure.
On the pleurites, — each pleurite bears four very powerful bristles,
one opposite the anterior annulus, one immediately ventrad of the
6 Journal of Entomology and Zoologj'
rudimentary spiracle, the remaining two in a transverse row on
the posterior ring near the caudal margin ; on the eighth pleurite
there is a single bristle. On the sternites, no bristles. Terebra
of the ovipositor ending almost on a common level, the tergal valves
a very little longer; each tergal valve terminates in four rather
weak bristles. In the male pupae, the sternal valves are slightly
more tumid and project beyond the level of the tergal valves.
Nepionotype. Urbana, Illinois. September 19, 1920.
Neanotype. Urbana, Illinois, September 29, 1920.
Paratypes, larvae and pupae, September 5, 19, 29, 1920.
Fig
Fig
Fig.
Fig.
Fig,
Fig,
Fig,
Fig.
EXPLANATION OF PLATE
Head capsule of larva, ventral aspect.
Mandible
Apex of mental bai'.
Antenna of larva.
Spiracular disk of larva, dorsal a.spcct.
Spiracular disk of larva, lateral aspect.
Pupa, lateral aspect.
Head of pupa, ventral aspect.
Ant.= Antenna; Lb. = Labium; M.\. = Maxilla; P.= Maxillary sheaths
A New Platydesmoid Diplopod from
California
By Ralph V. Chamberlin
From Dr. Hilton I have received an adult and several imma-
ture specimens of the interesting new genus below described.
The male is not yet known.
GosodesmiLs, gen. nov.
A genus differing from Platydesmus and Brachycybe in its
much narrower keels, the body as a whole being slender, more as in
Dolistenus and Pseudodesmus, body differing from that of the last
mentioned genus in being much more depressed, the keels hori-
zontal or, on anterior segments, upturned. Keels for the most part
laterally a little thickened or margined. Dorsum of each segment
with two transverse rows of large tubercles which are laterally
compressed, in part cariniform, the median ones not greatly en-
larged as in Pseudodesmus, pores not pedicillate; opening at
margin. Fifth segment normal. Head as in Brachycybe : no eyes
present.
Genotype. — G. claremontiis, sp. nov.
Gosodesmus claremontus, sp. nov.
The dorsum of the type is fulvus, in part of a distinct reddish
or pink tinge. The venter paler.
Head shaped nearly as in Brachcybe lecontii but somewhat
narrower and the antennae a little more clavate.
Keels of first five segments bent forwards, laterally strongly
rounded. Keels of sixth and seventh segments also bent forwards
but with the lateral margins straight at middle, the corners, how-
ever, widely rounded. On subsequent segments the keels have the
posterior corners extended a little caudad, the production becoming
pronounced in the caudal region. Keels of the penult segment pro-
duced directly caudad, nearly as far as caudal margin of last
tergite. Lateral margins of keels caudad of the eighth with
straight portion longer, slightly indented at middle, margined.
The angles on all keels remain rounded, but the posterior ones in
the more caudal segments narrowly so. Caudal margin of keels
toward mesal or proximal end bulging or shouldered, the caudally
extending portion abutting against or a little overlapping the
anterior border of the succeeding keel. First tergite with six
tubercles in each row, or with one or two extra ones in an indistinct
third row along anterior border. Tergites of middle region of
10
Journal of F,nt()iii<>li)t;\ and Zoology
body with mostly ten tubercles in the anterior row. and six or eight
in the posterior one.
Anal tergute broad, sides straight, caudal margin gently
convex.
Number of segments in type (female), fifty-two.
Length, 13 mm.; width, 1.2-mm.
Locality. — California : Claremont.
^\p±kh>
Gosodesmiis claremont us, sp. nov. Anterior view of head to
left above, below dorsal view of seventeenth tergite. On the right,
dorsal view of head and first four tergites, with right antenna
omitted. x55.
Hydroids Near Laguna Beach
Arthur S. Campbell
The hydroid fauna of Laguna Beach has been little studied but
there are a number of interesting forms to be found there. A few
collections, made almost at random from time to time, and with
no special search, form the basis of these notes.
The excellent papers by Torrey, Calkins, and Nutting have been
freely consulted in making the determinations. The splendid mon-
ograph by C. C. Nutting is especially invaluable to all who may have
to do with a systematic discussion of the group.
The more valuable results of this short paper are the distribu-
tional and systematic records, together with notes concerning
ecological and breeding relations. More extensive studies will re-
veal much data not hitherto brought to light concerning the
ecology, life-histories, variations, and other bionomical details of
the group in this interesting locality.
Key to the Hydroids of Laguna Beach
A. Hydranth without hydrotheca.
B. Hydranth with a basal whorl of filiform tentacles.
C. Hydranths solitary. Large. Corymorpha palma.
CC. Hydranths colonial.
D. Branched profusely. Medium size, often pinkish
in color.
Tubiilaria crocea.
DD. Branched sparsely.
Tubular ia sp.
BB. Hydranth with distal, knobbed tentacles.
Syncoryne mirabilis.
AA. Hydranth with hydrotheca.
B. Hydrotheca sessile. Gonangia are sporosacs.
C. Hydrotheca in two rows, usually opposite, on the stem.
D. Hydrotheca margin with two teeth.
Sei'tularia furcatn.
DD. Hydrotheca margin smooth, tubular, adnate at
base.
Sertularia tricuspidata.
12 Journal of Entrjmologj- and Zoology
CC. Hydrotheca in a single row on stem.
D. Hydrocladia on erect stems.
E. One or more intermediate internodes. Hydro-
theca as deep as long.
PI u malaria setacea.
EE. Septal ridges moderate, usually two in each in-
ternode.
Plumularia lageniffra.
DD. Hydrocladia modified as corbulae protecting gono-
theca.
E. Median tooth straight. Nine teeth.
Aglaophenia pluma
EE. Eleven teeth, irregular.
Aglaophenia !>trntlii(inide^
BB. Hydrotheca stalked, bell-shaped.
C. Gonophores are sporosacs.
CC. Gonophores are medusae.
D. Pedicels in pairs.
Ca ni pa n ula ria c.rig iin .
Obelia gracilis.
DD. Pedicels not in pairs.
E. Pedicels on shoulders produced from stem.
Obelia geniculata.
EE. Pedicels not geniculated, branching on all
sides.
Obelia commissurolis.
Sertnlaria de.tmoide'^ Torrey and Eitdendriiini ramomni L.
obtained during the summer of 102' are not included in this key.
They were determined for us by Mr. W. S. Wallace of the Hopkins
Marine Station, Pacific Grove, Calif.
TlBlLARIAE
Corynidae: No basal whorl of tentacles, but with tentacles
scattered irregularly over the hydranth. Tentacles knobbed. Hy-
droid branched.
Syncorync mirabilix (Ag. ) Torrey. U. C. pul). Zool. Vol. 1.
1902. p. 31.
Hydranth cylindrical. Proboscis conical. Scattering, capitate
tentacles. Small. Bathymetrical range; exposed to breakers of
open sea or in quiet harbours, ours on exposed pier with 0. com-
Pomuna CoUejic Clart'iiioiit. California 13
missuralis, on live Mytilus. Abundant. With medusae in Decem-
ber, 1920.
CoRYMORPHiDAE : Large, solitary hydranth with basal and
distal whorls of filiform tentacles. Medusae produced just within
basal tentacles.
Conjmorpha palma Torrey. Hyd. Pacific Coast. U. C. pub.
Zool. vol. 1, no. 1, p. 37.
A very large and beautiful species found abundantly in quiet
pools. Solitary, rooted in sand by filamentous processes. Proximal
tentacles 18-30 in number. Balboa Bay, in sandy pool. Usually
numerous in unexposed places.
Tubulariidae: Solitary or colonial. Large, often bright
pink in color. Hydranths with a basal and a distal whorl of filiform
tentacles. Sporosacs are pendant clusters.
Tubularia crocea (Ag.) Allman. Gym. Hyds. 1871. Dense col-
onies, 8-10 cms. i"n length. Few branches. About 20-24 basal ten-
tacles. On piles with other hydroids. tunicates, Crustacea and
mollusca. Low tide, December, 1920. Long Beach, Gal.
Tubularia sp. Distinguishable from above species in several
characters but not corresponding with any available discriptions.
I am not inclined to think it the T. marina of Torrey. Growing
with the above species at Long Beach. Rather rare. Probably the
same species discussed by Professor Bean in the Fourth Laguna
Report of Pomona College. Specimens also collected during the
summer of 1921.
Sertulariidae : Colony usually branching; hydrothecae ses-
sile, forming a double row along opposite side,> of hydrocaulus;
gonangia large, few, no free medusae.
Sertularia furcata Trask. Proc. Calif. Acad. Sc, 1854, I, p.
112.
This is a very variable species but ours are typical and agree
with figured specimens of several authors. Gonangia were numer-
ous on colonies taken at Huntington Beach, April, 1921, from piles
under the pier. Numerous, on stalks of algae and on rope.
Sertularia tricKspidata Hincks. Hist. Brit. Zoophytes. London,
1868.
This is a very common species at Laguna Beach, .growing in
great numbers on Fucus with other hydroids. Inshore tide zone.
January, 1921. With a creeping rootstock on which there are a few
gonangia, ripe. Hilton.
14 journal nt Knt<minliifjy and Zoology
Campanulariae
Plumulariidae : Hydranths t^essile, borne in a row on small
lateral branches, with nematophores. Gonangia large.
Plumularia fietacea (Ellis) Lamark. Anim. sans Vert.. 1st
ed. 1815. p. 129. Large. Nematophores, 2 above and one below
hydranth. Alternate hydrocladia. Not branched.
On piles under the Pleasure Pier, Long Beach. Calif. Low tide.
December 1920. Gonangia ripe, in pairs.
Plumularia lagenifera Allman. Jour. Linn. Soc. Lond., 1885,
XXIX. p. 157, pi. XXVL Very large and stiff. Unbranched. Cor-
i)ula not numerous. Station unknown, probably from dredgings at
Laguna Beach by Bean.
Aylaophcnia ^truthionides (Murry) Clark. Trans. Conn. Acad.,
Ill, 1876, p. 272. Small, abundant on Fucus inshore, at Laguna
Beach. With ripe gonangia January, 1921. Hilton. Commonest
hydroid.
Agkiophenia plitma (Linn.) Lamx., Hist. Pol. F'lex. 1816.
Some very typical specimens of this species were taken from
near the end of the pier at Huntington Beach, California. Readily
distinguishable by striking contra.st in color of dark .stem and light-
er hydrocladia. Rather tall. With corbulae in April, 1921.
EUCOPIDAE: Colonial, either branched or simple; hydrothecae
campanulate, stalked; aperture toothed or not; gonangium large
usually in axil of branch, free medusae.
ObcUa comniissuralis McCr. Gym. Charls. Harb., p. 95.
High, sparsely branched colonies. Hydranth deeply campan-
ulate. Pedicels annulate throughout, alternate.
On live Mytilus with other hydroids. Long Beach Pier, not
rare. December, 1920.
OhcUa qcnicultttn (Linn.) Schulzc. Nordsee Exped. 1872. j).
129.
An abundant shoi-e form everywhere. On Funis inshore. La-
guna Beach, Calif. January, 1921. No gonangia.
Ohelia gracilis Calkins. Some Hvdroids of Puget Sound, 1899,
p. 353.
Some typical specimens were taken on stems of other hydroids
from pier at Huntington Beach, California. No gonangia in April,
1921.
Pomona College, Claremont. California 15
Campakulariidae: Either a branched or a simple colony on
which are campanulate and usually stalked hydrothecae; hypos-
tome trumpet-shaped. Gonangium large, never with free medusae.
Campanularia e.rigua (Sars) Van Beneden. Rec. sur la Faune
Littorale de Belgique. 1867, p. 163.
This species I am not at all certain about but my specimens
seem to agree very well with descriptions and keys of both Nutting
and Torrey. If the identification should prove correct the species
has a much more southerly distribution than hitherto reported.
Calkins reports it from Puget Sound but Torrey has not included
it in his descriptions. The species is decidedly northern.
The gonangia of my specimens are a little fuller and with more
pronounced opercula than that figured by Nutting, but as those of
my specimens are ripe and those figured by Nutting are not this
may be of little significance.
Specimens on thalli of a seaweed, probably Macrocystis, exact
source unknown. Bottle labeled. Illingsworth, Pacific Grove,
July, 1899. Many ripe gonangia.
(Contribution from the Laguna Marine Laboratorv of Pomona
College.)
VIII.
Rotifera, Gastrotricha and Kinorhyncha
Rotifers. The usual type of nervous system of the female is
a dorsal ganglion or brain from which slender nerves pass to ten-
tacles, the ciliary disc or the general body. In Disco-pus, Zelinka
shows a ventral oesophageal ganglion as well as the usual dorsal
one. The shape of the brain or dorsal ganglion differs somewhat
in various species being almost spherical in some and quite elongate
in others, and in many cases bi-lobed. In Frullaria several longi-
tudinal strands of the nervous system have been shown. At nodal
points ganglion cells are located. The peripheral nerves are chiefly
as follows : one to each of the tentacles ; a pair of lateral nerves
which descend into the body and divide into two main branches,
one more ventral and one lateral, which give off numerous lateral
divisions to the muscles and viscera. Many fine branches run from
the brain to the ciliary ring to end in intimate relation with the
ciliary cells.
Antennae or feelers, usually three in number, a median dorsal
and two lateral ones are supplied by definite nerves. Each of these
structures consists of a small cluster of sense hairs born on a
slight swelling which receives the nerve. Sometimes the antennae
are retractile by means of internal muscle. These antennae may be
organs for touch or smell or both.
The brains of some forms contain a sac full of mineral mate-
rial. This may be some sort of sense organ, possibly a statocyst.
The eye-spot in its simplest form is a refractive globule in a
red pigmented cup to which latter nerve fibers pass, or the eye or
eyes may rest directly on the brain. Sometimes two eyes occur and
these may be very close together, almost like one. In some species
the eyes are just under the ciliary band or within the disc. A me-
dian and two lateral eyes occur in some forms, or even another pair
of eyes may also be found. In some, pigment spots occur at the
hind end of the body.
It has been suggested that some rotifers are able to avoid ob-
jects by means of a sense of sight aided by the tactile and olfactory
sense.
The chief work on the nervous system of this group has been
by Zelinka, 1888-90, Cast, 1900, and Halva, 1905. The more recent
work of Hirschfelder, 1910, has added quite a little to our knowl-
edge of the nervous system and sense organs in a number of forms.
This last author recognizes four general types of nerve cells which
grade into each other to some degree. Nerve fibers are described
as containing a central core of fibrils, an intermediate covering and
an outer sheath. Cells ai-e uni- or bi-polar; the last kind has one
IS
Joiirniil of Entiinv>l(i^:y and Znoloj:)
process passing to the periphery, the other running centrally. The
number, position, size and form of the cells is symmetrical in both
halves of the ganglion, also the processes are symmetrically dis-
posed. Commissural fibers bind right and left halves of the
ganglion. Some fibers leave the brain directly from ganglion cells
while others enter or leave by way of paired nerve fibers which
connect directly with the central fibrous core of the ganglion.
The ganglion cells are said to be al)solutely constant as to their
position, form and relative size. The position of the nuclei in the
cytoplasm is not so constant. The larger and smaller fibers seem
also constant in number and position.
Sense cells are found at the surface of the body more or less
removed from the surface. Single sensory nerve cells with two
nuclei end directly in the surface. Another kind of sensory ending
A. Rhizotidd, front and profile, showinj; position of the nervous system.
Zelinka.
B. Embryonic stage of a rotifer showing position of nervous system in
two dark masses. Zelinka.
C. Fnillaria, showing position of nerve strands in the body.
D. Position of nervous system in rotifer after Pelage et Herouard.
E. Nervous system of Floscularia after Hudson and Gosse.
F. DincopiiH, showing nervous system, after Zelinka.
G. Sense organs with nerves from the brain shown in cross section
after D. & H.
H. Nervous system of Echinoderes from several sources.
I. Nervous system of Echinoderes from above. Schepotieff, but much
changed.
J. K. Head end of Chaeltnintun from helow J. and above K. showing brain
and sense hairs. Zelinka.
Pomona Colleije, Clarcmont, California I''
is found in the tentacles where there is a combination of sensory
cells at the base of the sense organ.
The retrocerebral apparatus in Eosphora consists of two glands
lying back of the brain. They are covered with membrane and so
not in direct connection with the brain. One of these glands is the
pear-shaped retrocerebral sac which is clear with vacuoles. If
this is in any way a sense organ it is a question what its function
would be.
In Eosphora there is a single eye on the surface of the brain
and two slightly pigmented knobs at the anterior margin of the
animal ; these have a direct connection with the brain and must be
sense organs, possibly something like eye spots.
Gastrotricha. In 1864 Gosse described a knob on the oesoph-
agus as the brain in Chaetonotus. Ludwig in 1875 described the
nervous system on the side of the brain. Butschli, 1876, added
nothing of importance and Fernald, 1890, did not see the brain in
Chaetonotits. The clearest recognition of the nervous system was
by Zelinka in 1890. A large brain in the head region surrounds
the gullet above and on the sides and a pair of nerve trunks extend
down the body. Cephalic sense hairs are directly connected with
nerve cells of the brain. The hairs of the body may be for touch
or possibly smell or taste. Simple eyes have been described for a
number of species in the back part of the head, as small red spots,
but not all species possess them.
KiNORHYNCHA. Claparede in 1863 describes a nervous sys-
tem in this group and others at an early time also figure or describe
something of the nervous system. Reinhard, 1887, believes that in
most cases the nervous system was not seen by the earlier investi-
gators. He describes and figures a ganglion on the oesophagus but
gives no details. Zelinka, 1894, describes a circum-oral ring and a
long ventral nerve strand. Schepotieff, 1907, describes a brain
above the oesophagus with two connectives and a ventral strand.
The nervous system is somewhat like that of Gastrotrichia with a
large upper brain of a large mass of three general parts all fused.
The ventral strand runs the length of the body but is not differen-
tiated into ganglia but has cells along its course. Eye spots have
been described in some species, the number being from 2-8.
BIBLIOGRAPHY
Butschli, 0.
1S7(). Untersuchungen uber freilebende hematoden und die Gattung
Chaetonotus. Zeit. f. wiss. Zoll. Bd. 26. pp. 363-413 Taf. 23-26.
Claparede, E.
1863. Beobachtungen ubei- Anatomie und Entwickelungsgeschichte wir-
belloser Thiere. An. der Kuste Normende pp. 90-92, Taf. 16, figs.
7-16.
20 Journ.ll of Entnmolrjjjy and Zool()f:\
Eckstein, K.
1883. Die Rotatorien der Umgegend von Gissen. Zeit. f. wiss. Zool. Bd.
39, pp. 333-433, Taf. 23-28.
Fernald, C. H.
1883. Notes on Chaetonotus larus. Am. Nat. Vol. 17, pt. ii, No. 7.
Cast, R.
1900. Beitrage zur Kenntnis von Aspsilus vorax Leidy. Zeit. f. wiss.
Zoll. Bd. 67, pp. 167-214.
Gosse, P. H.
1864. The Natural History of the Hairy-backed Animalcules (Chaeton-
otidae). The Intellectual Observer, vol. .'>. pp. 387-406. pi. 1-2.
Halva, S.
1905. Beitrage zur Kenntnis der Radertiere I. Zeit. f. wiss. Zool. Bd.
80, pp. 282, 321. Taf. 17-18.
Hirschfelder, G.
1910. Beitrage zur Histologie der Radertiere. Zeit. f. wiss. Zool. Bd.
96, pp. 209-335, Taf. 9-13, 9 text. fig.
Ludwig, H.
1875. Ueber die Ordung Gastrotricha. Zeit. f. wiss. Zool. Bd. 26. pp.
193-226, Taf. 14.
Mobius, H.
1875. Ein Beitrage zur Anatomie des Branchionus plecatilis Mull. Zeit.
f. wiss. Zool. Bd. 25, pp. 103-113. T.if. 5.
Reinhard, W.
If87. Kinorhyncha, ihr Anatomischer Bau und ihre stellung ini system.
Zeit. f. wiss. Zool. Bd. 45, pp. 401-467. Taf. 20-22.
Schepotieff, A.
1907. Die Echinododeriden. Zeit. f. wiss. Zool. Bd. 88, pp. 291-326. Taf.
17-20.
Stokes, A. C.
1887. Observations on Chaetonotus. Jour. d. Microgr. v. .\i, pp. 77-85,
150-153, 566-565. v. xii, pp. 19-22, 49-51.
Sarasin, P. and F. Sarasin.
1888. Ueber die Anatomie der Echenothuriden und Phylogenie der Echen-
odermen. Krgeb. Naturn. Forseh. auf. Ceylon, pp. 84-154, Taf.
X-XVII.
Schultze, M.
1853. Ueber Chaetonotus, etc. .'\rch. f. .\njit. u. Phys. Bd. 6, pp. 241-254.
Wicrzcjski. A.
1893. Atrochus tentaculatus nov. gen. et. sp. Zeit. f. wiss. Zool. Bil.
55, pp. 696-712.
Zclinka, C.
1888. Studien ueber Raderthiere. Zeit. f. Wiss. Zool. Bd. 47, pp. 353-458.
Taf. 30-34. 4 wood cuts.
l!-90. Die Gastrotrichen. Zeit. f. wiss. Zool. Bd. 49, pp. 209-384. Taf.
11-15, 10 wood cuts.
1894. Ueber die Organisation von Echinoderes. Verb. deut. Zool. Gesell.
Bd. 4, pp. 46-49.
I& NOV 17 1939 «:
VOLUME FOURTEEN NUMBER TWO
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
JUNE, 1922
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT of ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
A Catalog of the California Aleyrodidaf and the Description
OF Four New Species — Donald D. Penny 21
Preliminary Notes on the Growth-staces in Brittle-stars —
Arthur S. Campbell 37
The Nervous System and Sense Organs, IX — //'. A. Hilton 45
Kutered Clwejnoiit, Cal.. Foit-Office Oct. 1, isio. as secocd-ulass njaller, under Act of Congress ol
Marcli 3. 187«
Journal of Entomology and Zoology — Advertising Section
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A Catalog of the California Aleyrodidae
and the Descriptions of Four
New Species
By Donald D. Penny
Introduction
This paper consists of a list of the ah-eady described species of
Aleyrodidae, or white flies, taken from the State of California, and
a record of their food plants and localities together with the descrip-
tions of four new species.
The writer has not attempted to give a systematic arrange-
ment of the family in this paper but has laid much stress on the
completeness of the list of food plants upon which the different
species have been taken in order that from a knowledge of the food
plants the family will be more readily accessible and at the same
time may be kept up to date in respect to the host records.
In the collecting of specimens the writer has not been con-
fined to any one section of the state but has taken and received
specimens from a wide range of localities, including sections of
both high and low elevation, from the extreme north to the extreme
south of the state. This has resulted in the recording of some new
hosts for the already described species as well as the finding of
the four new species herein described.
The writer desires to thank Professor E. 0. Essig for the
many specimens given and other kind assistance rendered during
the preparation of this paper.
Paratypes of the author's new species have been deposited
with the collection of the California Academy of Sciences, Golden
Gate Park, San Francisco, California.
Dialeiirodf:^ cit7-i (Riley and Howard)
(Alojrodcs citri Riley and Howard)
Syn. : aurantii Maskell
1893— Insect Life, vol. .5, p. 219.
Food Plants. — Ailantlius glandulosa, AUamanda neriifolia.
Ampelopsis tricxspidata, Cera ic^ sp., Choisya ternafa. Citrus spp.,
Coffea arabica. Diospyros kaki, Diospyros tnrginiana, Ficus macro-
phylln. Frnxinm lanccoJata, Gardenia florida. Gardenia ja-imin-
oides, Hedera helix, Jasminum fruticans, Jasminum odoratissimum .
Ligustrum amurense, Ligustrum sp., Madura aurantiaca, Melia
azedarach. Melia azedarach var. umbra culif or mis, Myrtus com-
munis. Magnolia fuscata. Myrtus lagerstroemia, Osmanthus amer-
22 Journal of Kinomolouy :ind Zoolo[:y
icaniui, Prunus caroliniana, Pntnia laurncerasiis, Punica {/ranaium.
Pyrns sp., Quercus aquaticn. Sniihi.r sp., Syritiga I'ulgaris, Tecomo
radicana. Viburnum thiits, Xa)itho.ryIu»i clai'aherculis.
Localities. — At present this species is known to exist in the
cities of Sacramento and Marysville. Also occurs in the Southern
States — North Carolina, South Carolina, Georgia, Florida. Ala-
bama, Mississippi, Louisiana and Texas.
Dialeurodes citrifoUi (Morgan)
(Aleyrorhs citrifoUi Morgan)
Syn. : nuhifera Berger
1910— E. W. Berger-Rull. 103, Fla. Agr. Exp. Sta. (A.
nubifera).
Food Plants. — Citrus spp.
Localities. — Not in California at the present time, having
been once exterminated at Bakersfield. Also recorded from Miss-
issippi. North Carolina. Louisiana. Florida. Cuba, China. Japan
and India.
Aleurophiti('< coronaiw^ (Quaintance)
( Alcitrodci (oronata Quaintance)
1900— A. L. Quaintance, Tech. Ser. No. 8, Div. Entom. U. S.
D. A., pp. 22-23. Orig. desc.
Food Plants. — Arhutw nunziesii, CaManea sp., Hetcromeles
arbutifolia, Quercus agrifolia. Qucrcu< chrysolepif, Quercus dcwi-
flora. Collected by the writer on Rhamnun califnrnica at Collins
Springs, May 1917.
Localities. — Alameda County. Collins Springs, Golden Gate
Park, King's Mountain. Lns Angoios, Mendocino County. Pomona.
San Bernardino, Santa Catalina Islands, Santa Clara Valley, Santa
Cruz Range, San Ramon Valley, Santa Rosa, San Jacinto, Sierra
Morena Range, Yo.semite Valley.
Aleuroplatus f/ehfitiosus (Cockerell)
( Aleyrndc.< <idatino<ii-< Cockerell)
1898— T. D. A. Cockerell, Can. Entom. Vol. 30. p. 264. Orig.
desc.
Food Plants. — Qucrcwi ofirifoUa. Quercus nrizonica. Col-
lected by the writer on Rhnmtius caUfnrnica at Collins Springs,
May 1917.
Localities. — Collins Sjirings. Collected by E. 0. E.ssig at
Auburn and Placerville. Also occurs in Arizona (type locality).
Pomona College, Clareniont, Calitornia 23
Pealius kelloggi (Bemis)
(Aleyrodes kelloggi Bemis)
1904 — Bemis. Proc. U. S. Nat. Mus., Vol. 27, p. 499. Orig.
desc.
Food Plants. — Prunus ilicifolia, Quercus agrifolia. Collected
by E. 0. Essig on Catalina cherry, Pasadena, Dec. 1914. Also
Niles.
Localities. — Niles, Pasadena, Santa Clara County, Sierra
Morena Range.
Pealitis maskelli (Bemis)
(Aleyrodes maskelli Bemis)
1904.— Proc. U. S. Nat. Mus., vol. 27, p. 524. Orig desc.
Food Plant. — Quercus densi flora.
Localities. — King's Mountain, La Honda.
Bemisia, inconspicva (Quaintance)
(Aleurodes incovspicua Quaintance)
1900— A. L. Quaintance. Tech. Ser. No. 8, Div. Ent. U. S.
D. A., pp. 28-29. Orig. desc.
Food Plants. — A7-butHS menziesii, Clematis ligtistici folia.
Heteromeles arbutifolia, okra, Physalis sp., Quercus agrifolia.
Quercus densiflora, Rhamnus californica, Rhamnus crocea, sweet
potato, Umbellularia californica. Collected by the writer on Acer
macrophyllum. Los Gatos, October 1916.
Localities. — Haywards, Los Gatos, Santa Cruz and Sierra
Morena mountains. Also recorded by Quaintance from Florida.
Aleyrodes amnicola Bemis
1904— Proc. U. S. Nat. Mus., vol. 27, p. 514. Orig. desc.
Food Plants. — Sali.i- laevigata, Washingtonia nuda.
Localities. — Stevens Creek.
Aleyrodes essigi. new species
Larva : — Color pale yellow with orange colored visceral glands
in abdominal region ; shape flat, elliptical, broadest in the abdom-
inal region. The dorsum is free from wax but there is a lateral
fringe of coalesced, white, wax rods. Segments of the case dis-
tinct. Marginal crenulations well rounded and incisions deep.
Cephalic margin with a pair of short, delicate setae; caudal margin
with long caudal setae set in tubercled bases ; latero-caudal margins
with a pair of small setae and a similar pair ju.st within the latero-
caudal margins.
24
Journal of FCntomology and Zoology
Pupa-case: — Size 0.92 mm. by 0.53 mm.; shape elliptical;
color transparent white with developing adult within light yellow.
There is a vertical ventral fringe of closely coalesced, white, wax
rods e.xtending to the leaf. This fringe may be seen adhering to
the leaf in the form of a ring when the ca.se is removed. Secre-
tions of the dorsum are lacking. The segments of the dorsum are
distinct. The abdomen bears two parallel rows of irregular
depressions which are indistinct in some cases. Vasiform orifice
subsemielliptical with the cephalic margin .straight. Operculum
subrectangular, curved laterally, the distal end truncate extending
one-half the length of the orifice. Lingula cylindrical extending
about seven-eighths of the length of the orifice, densely setose at its
distal end which bears a pair of small lobes and a pair of long,
straight setae projecting caudad. There are other setae on the
V\g. \. Alci/rodcn esgigi n. sp. a, pupa-case; b, vasiform orifice; c, anterior
margin of the forewinp; d, forewinp; c, claw of the adult; f, antenna
of the adult.
case as follows: a small, delicate
long jjair in the cephalic region,
abdominal segment, a minute i)aii
of the cephalic margin of the va
tubercled bases just within the c
pair on the latero-caudal margi
carefully in mounting to jircvent
marginal area is not set ofi" from
depression. The margin of the
pair on the cephalic margin, a
in e(iually long jiair on the first
r opposite the lateral extremities
iform orifice, a long pair set in
audal margin and a very small
n. The case must be handled
breaidng these setae. The sub-
the dorsum by a raised ridge or
case is evenly crenulated. the
Pomona Collejic, Clarciiiont, California 25
incisions shallow except at the caudal margin where they become
deeper and the wax tubes longer and narrower. The thoracic
tracheal folds are not evident.
Adult female: — Length 1.2 "mm.; general color light yellow
with head and thorax darker than abdomen. Legs and mentum
dusky. Paronychium blade-like ; wings immaculate. Forewing
length 1.2 mm., width 0.40 mm. Radial sector, media and cubitus
present : radial sector with two flexures. Media short ; cubitus a
faint line except at the base. Antennae dusky ; average lengths
of segments as follows: segment 1, 0.024 mm., segment 2, 0.158
mm., segment o, 0.109 mm., segment 4. 0.062 mm., segment 5, 0.072,
segment 6, 0.044 mm., segment 7, 0.056 mm. Eyes very dark red,
constricted but not divided.
This species was collected by Professor E. 0. Essig, for whom
the species is named, on Ulmus sp. at Mission San Jose, September
1916.
Aleyrodes pruinosa Bemis
1904— Proc. U. S. Nat. Mus., vol. 27, p. 491.
Food Plant. — Heteromeles arbutifolia.
Localities. — Berkeley, Catalina Islands, Leland Stanford Jr.
University.
Aleyrodes spiraeoides Quaintance
1900— A. L. Quaintance. Tech. Ser. No. 8, Div. Entom. U. S.
D. A., pp. 36-38. Orig. desc.
Food Plants. — Aesculus calif ornica. Convolvulus sepium, Loni-
cera involucrata, Nicotiana glauca. Opulaster capitatus. Plantago
major, Sonrluis oleraccu=:. Solaniim douf/la^ii, Tro.riinon sp. Col-
lected by the writer on Asel'ipias sp., at Berkeley, October 1916
and in Santa Cruz County, November 1920; Ceanothus sp.. Los
Gatos, December 1917; Hypericwin androsamum on the University
of California Campus, November 1916: Melaleuca hypericifolia on
the University of California Campus, November 1916 and on Pen-
tastamen at Capitola, December 1917.
Localities. — Alameda, Berkeley, Los Angeles, Los Gatos, Santa
Cruz County.
Ah'urotulus nvpln olepidJs (Quaintance)
Syn. : extraniens Bemis
1900— A. L. Quaintance. Tech. Ser. No. 8, Div. Entom. U. S.
D. A., pp. 29-30. Orig. desc.
Food Plants. — Acrostichum capense, nephrolepis.
Localities. — Conservatories of San Francisco. Type locality,
Pennsylvania.
26 Journal ot Entomology and Zoology
Alevrothrixtis interrogationis (Bemis)
{Aleyrodes interrogationis (Bemis)
1904— Bemis, Proc. U. S. Nat. Mus., vol. 27. p. 516.
Food Plant. — Ceanothns calif ornicus.
Localities. — Black and King's mountains, Pacific Congress
Springs.
Aleuroparadoxv.s iridescens (Bemis)
(Aleyroden iridescen-t Bemis)
1904— Bemis. Proc. U. S. Nat. Mus.. vol. 27. p. 487. Orig.
desc.
Food Plants. — Arctostaphylos manzanita, Heteromelea arbuti-
folia. Rhamnus californica, Rhamnii^ crocea, Umbelhdaria cali-
fornica. Collected bv the writer on Salvia sp., San Diego County.
May 1917.
Localities. — King's Mountain. Yosemite Valley, San Diego
County. San Gabriel Mountains, Santa Clara Valley, Santa Cruz
Mountains.
Asterochiton coroUis, new species
Pupa-case: — Size 0.90 mm. by 0.61 mm.; shape elliptical with
the caudal end truncate; color dark brown. The wax .secretion of
the dorsum, as observed from somewhat imperfect specimens, con-
sists of three separate systems as follows ; first, a continuous mar-
ginal fringe extending entirely around the case, the rods of which
are short and loosely .ioinod. projecting directly toward the leaf
to about one-half of the distance from the margin to the leaf.
Second, a series extending continuously around the case just within
the margin. The rods of this system are long, white and closely
coalesced at their bases and extend upward for the greater part
of their lengths then outward over the case, separating into ribbon-
like structures at their extremities. Third, a series of short, thick
rods arranged in groups which arise mesad of the second svstem
and which project toward the center of the case. In addition to
the dorsal wax the pupa case secretes a high vertical fringe of
wax on which the case rests. The submarginal area bears a row
of large, conical, papilla-like pores the bases of which are clo.se
together, the pores themselves measuring about 0.02 mm. in
length. These pores undoubtedly secrete the long ribbon-like wax
.structure. On the dorsum jtrnpcr are irregularly shaiied. con-
spicuous, pore-like oiicnings which are arranged in groui)s. the
outer margins of which conform to the general curve of the case.
These groups are found as follows: two in the cephalic region con-
sisting of fourteen pores each, two in the thoracic region containing
about twelve openings each and two in the abdominal region con-
taining twenty-four each. Scattered through these pores are numer-
ous very small circular pores also two pair of similar circular pores
Pomnna Collesie, Claremont. California
27
in the cephalic region close to the median line, two pair on each
segment of the thorax, two pair each on segments 1 and 2 of the
abdomen, one pair each on segment 3, 4, 5 and 6 and three pair on
segment 7 of the abdomen. In the submarginal area between the
papilla-like pores and the margin is a row of the small circular
pores and in addition to these some may be found with no apparent
regularity near the bases of the submarginal papilla pores. The
sutures of the case are distinct, the last three of the abdomen
strongly reflexed caudad. The crenulations of the margin are
broad and the incisions shallow. The submarginal area is not set
off by a raised ridge or depression. Vasiform orifice subcordate,
with the anterior margin straight, the lateral margins with corru-
gations or folds extending inward and downward ; operculum
shaped similar to that of the orifice with the caudal end slightly
truncate, one-half filling the orifice: lingula subspatulate, densely
setose, extending about three-fourths of the length of the orifice,
bearing at its distal extremity three pair of lateral lobes and a pair
of terminal lobes. Thoracic tracheal folds not evident. Just
laterad of the anterior margin of the vasiform orifice is a pair of
Fig. 2. Asterocltttvn corollis ii. sp. a, pupa-case; b, vasiform orifice, caudal
furrow and section of the caudal margin; e, pupa-case showing the
wax seci'etions.
28
Journal of Entomol(it;\ and Zoology
fine, delicate hairs set in circular bases. Delicate latero-caudal
marginal hairs are present but cephalo-marginal hairs and caudal
spines are lacking.
Adults unknown.
This species was described from three specimens of the pupa
taken by the writer on Arctostaphylos manzanita at Pine Hijls,
San Diego County, May 1917.
Locality. — Pine Hills, (type)
Aisterochiton diasemus (Bemis)
(Aleyrodes diasemus Bemis)
1904— Bemis. Proc. U. S. Nat. Mus. vol. 27, p. 516. Orig. desc.
Food Plants. — Ribes glxtinosnm, Sijmpboricarpos raceynosus.
Localities. — Alameda, King's Mountain, Leland Stanford Jun-
ior University, Menlo Park, San Francisquito Creek.
Asterochiton diminutis, new species
Pupa case : — Size 0.53 mm. by 0.33 mm., shape elliptical, very
convex, extending high above the leaf particularly in the cephalic
region the ventral surface of which is projected into a blunt point.
Color smoky white, parasitized specimens very dark brown. The
wax secretion of the dorsum consists of an irregular row of
tapering, glassy, white, waxen rods arising in the submarginal area
and which extend upward and outward over the margin of the
case and are about as long as one-third the width of the case ; also
Kip. ."i. Aslerochitdii tliiiiiinitis n. sp. a, pupa-case; b, side view of tlie pupa-
case; c, vasiform oiKice and section of the caudal maijrin; d, fore-
winjT of the adult; e, anterior niarj;in of the forcwinjr; f, claw of the
adult; g, first three segnn'iils of the antenna of the adult; h, male
genitalia.
Pomona College, Claremont. California 29
a pair of similar waxen rods arising in the cephalic region and
extending upward over the case, a pair each on two segments of
the thorax and on segments 3, 4, 5 and 6 of the abdomen. In addi-
tion to the dorsal secretion the case bears a continuous high, ver-
tical fringe of coalesced, white, wax rods which extend from the
margin of the case to the leaf. This wax functions as a support
for the case and it remains firmly attached to the leaf when the
case is removed. The pores which give rise to the dorsal wax
secretion are large, 0.012 mm. in length, conical in shape and are
arranged in a rather irregular submarginal row of about sixty in
number ; also a pair of similar pores in the cephalic region, a pair
each on the two segments of the thorax and a pair each on segments
3, 4, 5 and 6 of the abdomen. Between the bases of the submarginal
wax pores may be found very small circular pores, a row of similar
circular pores between these and the margin of the case, and along
the dorso-meson from the cephalic region to the vasiform orifice a
pair for each segment, though occasionally missing on some seg-
ments. Segments of the dorsum are distinct and in the abdominal
region the sutures are strongly bent caudad. Vasiform orifice sub-
cordate, length 0.06 mm., anterior margin straight, inner lateral
margins with corrugations extending downward. Operculum sub-
semielliptical, about one-half filling the orifice. Lingula slightly
hidden, subspatulate, densely setose, projecting slightly beyond the
orifice and bearing at its distal extremity a pair of terminal and
three pair of lateral lobes. Submarginal area not set ofl" from the
dorsum by a raised ridge or depression. Thoracic tracheal folds not
visible. Conspicuous caudal spines present, arising just within the
caudal margin, but cephalo-marginal and latero-caudal marginal
spines absent.
Adult female: — Length 1.0 mm., general color yellow to
orange, eyes very dark brown, constricted but not divided. Fore-
wing length ].15 mm., radial sector, media and cubitus present.
The radial sector is the main vein of the wing extending through
the central area. The media is reduced to a remnant, being very
short and faint and arising as a branch of the radial sector. The
cubitus appears as a cleared line arising independently of the radial
sector and projecting caudad toward the margin then paralleling it
for a very short distance before ending. The portion of the wing
through which the cubitus passes is very slightly dusky or unclear
thus making the cleared vein more distinct.* Length of antennae
segments from segment 1 to segment 7 inclusive, as follows : 0.024
mm., 0.052 mm., 0.128 mm., 0.048 mm., 0.064 mm., 0.044 mm.,
0.040 mm. Paronychium blade-like.
This species was described from an abundance of pupae and
several adults taken by the writer on tarweed (Chamaebatia
*This type of vein is spoken of by Bemis (Proceedings of the U. S. Nat
Museum, vol. 27, page 493) as a long, oblique, anal fold.
30 Journal (if Entomology and Zoology
foUohsa) at Placerville, May, 1918. The pupae occur on both sides
of the leaves but the very small size of the pupa case together with
the numerous minute leaflets on which the case rests make it
exceedingly difficult to observe with the naked eye.
Locality. — Placerville. (type)
Asterochiton glacialis (Bemis)
(Aleyrodes glaciali.'^ Bemis)
1904— Bemis. Proc. U. S. Nat. Mus., vol. 27. p. 518. Orig. desc.
Food Plants. — Ceanothus calif ornicti.'i. Clematis liguf^ticifolia.
Optdaster capitatus. Querent densiflora, Rhamiius califoniica.
Rubus vitifoliui^. Sytnphoricarpos racemosii.'i. Taken by the writer
on a Salvia hvbrid on the University of California campus. Novem-
ber 1916.
Localities. — Alameda, Berkeley, King's Mountain, Santa Clara
Valley, Santa Cruz and Santa Morena ranges.
Asterochiton hutchingsi (Bemis)
(Aleyrodes hutchingsi Bemis)
1904— Bemis. Proc. U. S. Nat. Mus.. vol. 27, p. 532. Orig desc.
Food Plant. — Arctostaphylos sp.
Locality. — Yosemite Valley.
Asterochiton madroni (Bemis)
(Aleyrodes madroni Bemis)
1904— Bemis. Proc. U. S. Nat. Mus., vol. 27, p. 507. Orig desc.
Food Plant. — Arbutus menziesii.
Localities. — King's Mountain. Collected by the writer at
Berkeley, Los Gatos, Santa Cruz County.
Asterochiton merlini (Bemis)
(Aleyrodes merlini (Bemis)
1904— Bemis. Proc. U. S. Nat Mus., vol. 27, p. 512. Orig. desc.
Food Plants. — Arbutus menziesii. Collected on Arctostaphylos
spp. l)y K. H. Davis in San Diego County and by the writer on
Arctostaphylos sp. at Colfax, April 1920.
Localities. — Auburn, Colfax, King's Mountain, Placerville.
San Diego County. (Throughout Sierra Nevada Mountains.)
Asterochiton tentacidatiis (Bemis)
(Aleyrodes tentaculatus Bemis)
1904 — Bemis. Proc. U. S. Nat. Mus. vol. 27. p. 494. Orig. desc.
Food Plants. — Cleynatis ligusticifolia, Lonicera involucrata.
Opulaster capitatus, Quercus agrifolia, Quercus densiflora. Rhus
diversabola.
Localitv. — Alameda.
Pomona College, Claremont. California 31
Asterochiton vaporariorum (Westwood)
(Aleyrodes raporarionnn Westwood)
Syn. : nicotianae Maskell
Syn. : papiUifer Maskell
Syn. : lecanioides Maskell
1856 — Westwood. Card. Chron., p. 8.52. Orig. desc.
Food Plants. — Ageratum, Aphelandra, Aster, bean, Begonia,
Bignonia, Capsicum, Chrysanthemum, Citrullus vulgarts, Coleus,
Cucumis melo, Cucumis sativus, Fragaria sp., Geranium, Gonolo-
bus, Lactuca sativa, Lantanu commara, Nicotiana, Oxalis, Pelar-
gonium Ptrsea gratissima. Primula vulgaris, Rosa sp., Rubwi,
Salvia splendcns. Solanum melongina, Solanum pseudo-capsicum,
Tecoma, Vitis. Collected by the writer on Abutilon sp., Santa Cruz
county, July 1917; on Aralia cordata, October 1916, Datura sanguin-
ata, November 1916, Hclianthus californicus, November 1916,
Eupatorium ruparinm, November 1916 on the University of Cali-
fornia Campus ; on Quercus kelloggi, Santa Cruz county, June 1919,
on RJiannius californica, Los Gatos, November 1916 and on Rhus
divcrsiloba, Santa Cruz county, September 1918.
Localities. — Berkeley, Los (Jatos, Santa Cruz County. Santa
Rosa.
Asterochiton vittatus (Quaintance)
(Aleurod.es vittata Quaintance)
1900— A. L. Quaintance. Tech. Ser. No. 8, Bur. Entom. U. S.
D. A., p. 42, Grig. desc.
Food Plant. — Chapparal.
Localities. — Claremont, Ontario, Pomona.
Asterochiton ivellmanae (Bemis)
(Aleyrodes ivellmanae Bemis)
1904— Bemis. Proc. U. S. Nat. Mus. vol. 27, p. 525. Orig. desc.
Food Plant. — Rhammis californica.
Localities. — Leland Stanford Junior University, Stevens Creek.
Tetraleurodes acaciae (Quaintance)
(Aleyrodes acaciae Quaintance)
1900— A. L. Quaintance, Tech. Ser. No. 8. Div. Entom. U. S.
D. A., pp. 19-20. Orig. desc.
Food Plants. — Acacia, Bensera microphylla, Rhamnus cali-
fornica.
Localities. — Fullerton, Los Angeles, Ontario. Also recorded
from Lower California and Mexico.
32 Journal ot Entomology and Zoology
Tetraleurodes dorseyi (Kirkaldy)
(Aleyrodes dorseyi Kirkaldy)
Syn. : qt(aintaticei Bemis
1907— Kirkaldy. Bui. 2 Div. Ent. Bd. Comni. Agr. & Forestry.
Hawaii, p 52.
Food Plant. — Rhamnus crocea.
Locality. — Stevens Creek.
Tetraleurodes crrans (Bemis)
(Aleyrodes errans Bemis)
1904— Bemis. Proc. U. S. Nat. Mus., vol. 27, p. 500. Orig. desc.
Food Plants. — Arbutus menziesii. Umbelhilaria californica.
Collected by the writer on Aesculus californica. University of Cali-
fornia Campus, September 1916.
Localities. — Berkeley, King's Mountain, Leland Stanford Jun-
ior University, San Ramon Creek, Santa Clara Valley, Santa Cruz
Mountains, Redwood creek. Usal.
Tetraleurodes herberti. new species
Pupa-case : — Average size 0.92 mm. by 0.64 mm. ; shape sub-
elliptical, slightly more pointed at the caudal end; color shining
black. The case is closely applied to the leaf. Dorsum keeled
along the median line with the raised area somewhat wider in the
cephalic and thoracic regions than in the abdominal region. The
segments are distinct with suture lines well defined, particularly
across the keeled area. Outlines of rudimentary legs on the
ventral surface plainly visible through the case. In the cephalic
region on both sides of the median line is a circular mark or
depression which is bounded by two markings arranged as arcs of
concentric circles. Caudad of these markings and close to the
median line on either side is another group of two subcircular,
clearly defined markings or i)ore-like openings, one latero-caudad
of the other. In the thoracic region is still another pair of tri-
angularly arranged groups of three irregularly outlined depres-
sions, a pair on each segment suture from the thoraco abdominal
suture to the vasiform orifice. In the cephalic and thoracic region
on either side of the median line is a row of about five small cir-
cular pores which passes just iaterad of the groups of markings in
those regions to and including the first al)dominal segment. Smaller
circular pores are present just caudad of the abdominal depres-
sions, one for each depression. A pair of similar pores are found
just cephalo-laterad of the anterior margin of the vasiform orifice,
another pair Iaterad and still another i)air just cephalad of the
anterior margin of the vasifoi'm orifice. Vasiform orifice sub-
ovate, surrounded by thickened integument and raised well above
the general level of the dorsum; operculum filling the opening.
PoniDtia C(
Clareniont, California
33
Lingula partially obscured, spatulate, distal end spherical and
densely setose. The submarginal area bears a continuous row of
conspicuous circular pores which are about 0.014 mm. in diameter
and which project somewhat from the case. The submarginal
area is set off from the dorsum proper by a raised ridge which is
continuous around the case save in the cephalic region. The cren-
ulations of the margin are broad and well rounded, the incisions
shallow. Marginal wax tubes project mesad about one-third the
width of the margin. Thoracic tracheal folds not evident. A pair
of fine, delicate caudal spines are present just within the caudal
margin, set in tubercled bases. On the suture lines extending
cephalad from the vasiform orifice is a pair of very fine delicate
hairs set in circular bases. Cephalo-lateral and caudo-lateral mar-
ginal spines are lacking.
Adult : — Length about 0.65 mm. ; color yellow with the head
and thorax lighter. Eyes dark red, constricted but not divided.
Wings immaculate, forewing with Radius 1, cubitus and media
present; media short, faint and poorly defined. Paronychium
blade-like. The antennae of all specimens were broken.
Described from material taken by F. W. Herbert, for whom
the species is named, at Pleasanton, Alameda County, October 1918,
Fig. 4. Tetraleurodes herheti n. sp. a, pupa-case; b, vasiform orifice.
.?4 JournnI of EntoniDlofi) and Zoology
on black locust. The specimens consisted of pupae with only three
adults which were badly damaged. The pupae were attached to
both sides of the leaf.
Localities. — Pleasanton. (type)
Tetraleurodes melanops (Cockerell)
( Aleyrodes melanops Cockerell)
1903— Cockerell. Bui. 67. Fla. Agr. Expt. Sta. p. 665.
Food Plant. — Quercus sp.
Localities. — Alpine Tavern, Mt. Lowe.
Tetraleurodes nigrans (Bemis)
( Aleyrodes nigrana Bemis)
1904— Bemis. Proc. U. S. Nat. Mus. vol. 27, p. 522.
Food Plants. — Arbutus menzH'sii. Arctostaphylos mauzanitn.
Ceanothu.s californicus. Clematis ligusticifolia, Eriodictyon cali-
fornicum, Hcteromelcs arbutifoUa. Loniccra involucrata. Primus
ilicifolia, Rhamnus califoruica. Symphoricarpos racemosus, Umbel-
lidarin rnlifnrnica. Collected bv the writer on Salvia sp.. Corona.
May 1917.
Localities. — Corona, Black and Kings Mountains, Pacific Con-
gress Springs, San Ramon Valley, Santa Clara Valley, Santa Cruz
Range, slopes of Sierra Morena Range, Stevens Creek.
Tetraleurodes perileuca (Cockerell)
( Aleyrodes pcrih]uv< Cockerell)
1903— Cockerell. Bui. 67. Fla. Agr. Exp. Sta., p. 664.
Food Plant. — Quercns sp.
Localities. — La Jolla.
Tetraleurodes splcndens (Bemis)
(Aleyrodes spletulcn-i Bemis)
1904— Bemis. Proc. U. S. Nat. Mus. vol. 27, p. 489.
Food Plants. — Rhnmnus californiea. Arctostaphylos sp.
Localities. — Leland Stanford Junior Universitv, Yosemite
Valley.
Tetraleurodes stanfordi (Bemis)
(Aleyrodes stanfordi Bemis)
1904— Bemis. Proc. U. S. Nat. Mus. vol. 27. p. 508.
Food Plants. — Quercus agrifolia, Quercus densiflora. Collected
by the writer on Rhamnus sp., Fresno, May 1917.
Pomona College, Clareinont, California 35
Localities. — Big River, Mendocino County, Black Mountain,
Fresno, King's Mountain, Santa Clara Valley.
References : — The writer has made use of all available works
dealing with the descriptions of the species of the family of Aleyro-
didae and especially the following:
"Contributions Toward a Monograph of the American
Aleurodidae" by A. L. Quaintance. Technical Series
No. 8, Bur. Entom. U. S. Dept. Agr., 1900.
"The Aleyrodids, or Mealy-winged Flies, of California, with
References to Other American Species," by Florence E.
Bemis. Proceedings of the United States National
Museum, vol. .37, pages 471-537, 1904.
"Classification of the Aleyrodidae," Part I and II, by Quain-
tance and Baker. Technical Series, No. 27, Bur.
Entom. U. S. Department of Agriculture, 1913, 1914.
"A Contribution to Our Knowledge of the White Flies of
the Subfamily Aleyrodinae (Aleyrodidae)," by Quain-
tance and Baker. Proceedings of the United States
National Museum, vol. 51, pages 335-445, 1917.
Preliminary Notes on Growth-Stages in
Brittle-Stars
Arthur S. Campbell
There are a number of conditions to account for our present
lack of a rational system of the brittle-stars. One of the principal
reasons why the group is so difficult to classify lies in the profound
ignorance of their growth-changes. The excellent systematic
work of Ljungman, Lutkin, Lyman, Koehler and the two Clarks
have brought some thousand species to attention but the real rela-
tionship of these as larger groups is yet quite unsolved. There have
been several attempts to rationalize the classification, one by Bell,
1892, and more recently by Matsumoto, 1915. Neither of these sys-
tems is thoroughly based upon phylogenetic history, and hence, can-
not be conclusive since the state of our present knowledge is such
as to forbid any sweeping generalizations.
Although the chief reason for our lack of a rational system in
the group is this lack of attention upon growth-stages, another
lies in the general disregard of palaeontological evidence, and a fur-
ther reason because attention has been focused upon larval, rather
than post-larval, stages.
Material heretofore studied in connection with this problem
of gi'owth-stages in the young of ophiurans numbers less than one
dozen species, all of which are Atlantic or West Indian forms. My
own observations were made upon seven species, the members of
five families. All are the members of the littoral fauna of Southern
California. Specimens were collected in all accessible habitats and
studied after preservation.
The excellent plates for this paper are the work of Miss E.
Keyes, a student in Pomona College.
It is not always possible to tell just why one places a form in
this or that group for many characters are subtile and one is obliged
to depend very often upon general features. Especially is one
dependent upon as complete a series as possible in placing a juve-
nile. H. L. Clark, in his paper on growth-changes in some brittle-
stars expresses the only formulation of the very important con-
tribution of R. T. Jackson to the study of juvenile brittle-stars that
I have seen. This law is a very real help in determining possible
relationships between specimens otherwise obscure or impossible
to differentiate. Briefly stated, we may say that, as applied to
these forms, the base of an arm of a young form corresponds ex-
ceedingly suggestively with the tip of an arm of an adult specimen
of the same species. However, the extent of localization varies
greatly in different species, as I have found. One needs much
.study to determine accurately the position of a given specimen.
38 Journnl of Kntomolony and Zoology
It is hardly possible, as I have pointed out, to formulate a gen-
eral system of the group hut among the groups examined the
Opiiioh'pidac and the Opliiocfimidai are noticeably separate and,
containing few local genera of well marked characters can readily
be separated both among themselves and from other families. Be-
tween the families Opiiinfliricidac and Amphiuridae I have found
many points of contact as I did also between some Ophiadcrmatidae
and the Amphiuridae. Beyond these generalizations I do not care
to advance any opinion.
Following are my results upon those species examined. Ex-
tremes of measurement and a few notes on certain of the more
obvious structural details are given. Other details can be made out
from study of the plates.
Opiiiocnjptiis maridnsii^ Clark. The smallest si)ecimen meas-
ured had its disc one mm. in diameter and with arms one and a half
mm. long. Young of this species differ from adults in few skeletal
details. The buccal fissures seem less marked and the arms rela-
tively longer in proportion to the disc.
Ophiodcrma panamen-i < Lutkin. The smallest specimen
measured two mm. across the disc and with arms eight mm. long.
Juveniles of this species resemble adults in many points but they
di'T(M- in others. The disc is set well apart from the arms. The
characteristic notches between the arms in the adults are absent.
The radial shields are scarcely marked. The branchial spines are
set almost at right angles to the arms. Not fi'Jrured.
Ophioplocus esmarki Lyman. The smallest spei-imen meas-
ured was one mm. across the disc and with arms eight mm. long.
Juveniles of 0. esmarki are always distinguishable by pinkish
bands crossing the arms. This species is especially interesting on
account of the schizogony that young specimens undergo.
Amphiodia harharae Lyman. The disc of the smallest speci-
men measured was three mm. and the arms twenty-eight mm. long.
These are always to be distinguished by the exceedingly long arms
at least ten times the diameter of the disc. Young seem to bear
many points in common with O. panamayi-iis.
Ophioiieris avmdafa Le Conte. Specimens of this species vary
from about one half mm. to two mm. in diameter. They undergo
schizogony in an unc(|ual jilane in certain cases. Like the adults
the arms have three flattened arm-spii'cs and with banded arms.
Ophi(ipt( ris pajiillosd Lyman. Measurements of the smallest
specimen in this species found were for the disc three mm. and for
the arms ten mm. These are distinguishable by the flat upper arm-
plates and coarse arm-spines, but thc^e are both characters that
vary even in one specimen.
Ophiothrix spicidata he Conte. I found liut few specimens of
this although the adults are abundant. The smallest specimen
measured one mm. in diameter. The reduction in the comparative
Pomona Collejje, ClaiciiKint, California 39
size of the radial shields is especially noticeable in a series of spec-
imens. One of the interesting features of this species is the great
range of color variation found. This is true both in young and
adults.
Conclusions :
1. A rational system in the brittle-stars is lacking. Such a
system may result in part at least, from a complete study of growth-
stages.
2. Jackson's law of localized stages repeating phylogenetic his-
tory seems to be well vindicated in this and other studies.
3. The groups examined seem to bear certain relationships to
each other, as indicated above.
Literature
Bell, F. J. Contribution to the Classification of the Ophiuroids.
1892. Pro. Zool. Soc. London, pp. 175-183.
Clark, H. L. Growth-Changes in Brittle-Stars. 1914
Papers from the Tortugas Lab. of the Carnegie Inst. Was. vol.
5. pp. 93-125.
Jackson. R. T. Localized Stages in development in plants and ani-
mals. 1899. Bost. Soc. Nat. Hist., vol. 5, pp. 89-153.
Ludwig. H. Zur. Entwicklung.-geschichte de'^ Ophiurenskelettes.
1881. Zeit. f. w. Zool., vol. 34, pp. 333-365.
. Jugendformen von Ophiuren. 1899
Sit. d. K. Preuss. Akad. der Wiss. Berlin, vol. 14, pp. 210-200.
KXPLAKATION OF I'LATES
All lifiTurcs XIO
pinilata.
G; H, I, upper and lo\\er surfaci's of various sizes. K
I'iate I, Ophiolliiix
A. B; r, D; F
Dorsal view of one whose ventral side is like A.
Plate III. A, B; E, F, Amphipudia burhtirue; C, D, Amphipodia barbaraei
Plate II. A. and B, upper and lower surfaces of Ophiouermis a)niiil(il:i
C and D, upper and lower surfaces of six armed O. a7niulala.
E, F; G, H, upper and lower surfaces of Ophioplneiis esmarki.
I, J, upper and lower surfaces of Ophioptei-is papillosa.
Plate IV. A, B, upper and lower surfaces of O. anuulata. C, Dorsal, E, D,
I dorsal and ventral views of Ophi(-cr(/ptun maculosus.
L.
IX. The Brvozoa
ECTOPROCTA.
About the earliest observations on the nervous system of
these animals was by Dumortier and Van Beneden in 1843. They
described the central nervous system of fresh water forms as com-
posed of two ganglia above the oesophagus joined by commissures.
From the aboral part of the ganglion a pair of nerves runs to the
oesophagus. They also considered that a pair of nerves supplied
the epistome.
In 1848 Van Beneden speaks of but a single ganglion.
Allman, 1856, in fresh water forms describes a single unpaired
oval ganglion. The two oesophageal nerves are represented as an
oesophageal ring with enervation for the epistome.
Hyatt, 1865-1868, describes the central ganglion in Phimatella
with the ganglion concentrated. The two long arms of the animal
however, are capable of independent movement. The ganglion in
Trederecella is spindle-shaped. In Plumatella the ganglion is kid-
ney-shaped and as it doubles upon itself by movements of the
animal it becomes heart-shaped. He describes a true nerve ring
about the oesophagus. Nerves go to the middle and end intestine.
Hyatt also describes nerves to the epistome and to the tentacles.
Nitche, 1869-76, has studied bryozoans quite extensively. He
found a central cavity in the ganglion in embryonic stages. He
recognized an oesophageal ring, intestinal nerves, tentacle nerves.
He recognized on the tentacles bristles which he called taste bristles.
Claparede, 1871, in some bryozoans describes the nervous sys-
tem of colonial forms ; nerve strands running the length of the body
were recognized.
Kraepelin, 1887, found the center of the ganglion in adult
forms, and the shape of the ganglion of fresh water forms elipsoid.
He also recognized peripheral ganglion cells in the ganglion. Oral
nerves were seen, as well as nerves to the epistome.
Verworn, 1887, in a general way recognized ganglion cells.
Saefftiger, 1888, has especially added to our knowledge of the
distribution of the nerves to the tentacle crown ; he also considers a
sympathetic system but says nothing of the sense cells in the ten-
tacles although he describes the epithelium of parts of the animal.
Braem, 1890, describes the central ganglion of fresh water
forms as hollow with an outer thinner oesophageal and a ventral
thicker wall. He considers the inner part of the ganglion as largely
fibrous.
Oka, 1891, has considered fresh water forms, especially Pec-
tinatella. Like Saefftiger, he finds the ganglion with a cavity in
the mature state. The ganglion is compared to a spindle bent in
46 Journal of Entomology and Zoology
the form of a U, with the concavity fitted to the anal side of the
oesophagus in an oblique position with arms turned slightly up-
wards. The end of each makes a turn in the oral direction, and is
continuous with a large nerve trunk which goes to the lophophore
arm. The ganglion is in direct connection with the inner cell layer
of the oesophagus, the outer layer of the latter enveloping it on all
sides. The lojihophore nerve trunks are likewise located between
the outer and inner layers of the body-wall ; they run beneath the
outer layer of the lophophore covered below Ijy epithelium. The
ganglion contains a large cavity extending to the ends of the gan-
glion. The wall of the ventricle is very thin and epithelial in
nature on all sides but the bottom on the anal side, where it is very
thick as it joins the main part of the ganglion. This thick portion
is distinctly separated from the epithelial part and is well seen in
the fresh state as a somewhat reddish mass with a slight constric-
tion in the median i)lane of the iiolype. It is this jiart that Ilyatt
took for the ganglion which he described as composed of two lateral
masses connected by a thick commissure. The epithelial part is
hard to recognize in surface views. A cross section of the lop-
hophore trunks is kidney-shaped; in it the nerve cells are much
crowded; the nerve cells are spindle-shaped, bipolar, with nuclei
in the middle, closely packed together with few fibers between.
The nerve trunks are thick and large as compared with the ganglia.
The matter of a circum-oesophageal ring was not settled; this
author did not find it. The colonial nervous system found in some
marine Bryozoa for the purpo.se of controling the movements of
the members of the colony seem to be entirely lacking in the species
PectinateUa (nUtdnosa, and this fact agrees with the behavior of
the animals as they act independently.
Cori, 1898, does not give much further information about the
nervous system of bryozoans.
Delage and Herouard, 1897. in a numl>er of diagrams show the
liosition of the ganglion in marine ectoprocts as being a single small
ganglion ventral to the oesophagus. There are probal)!y nerves
going to the tentacles, to the body and to the alimentary canal, but
these are not clearly shown in any case. A ganglion in the avicu-
larium is shown by Delage and Herouard and they indicate by a
series of diagrams how this ganglion might have been derived from
a single zooid by a series of gradual transformations.
Ladewig, 1900, shows such a ganglion center in an aviculariuni
of a marine ectoproct.
The sensory system of ectoprocts has been described by
Nitsche on the tentacles of AlcifoncUa as stiff bristles to which he
ascribes the sense of taste. Verworu, Kraepelin, Braem and others
have seen these without ascribing special functions to them. It
seems probable that the tentacles must have some special sense
organs for touch or other senses.
Fig. 19. Nervous system of fresh water bryozoa. A. General view of the
nei'vous system of Cyistatella. B. Oi'al surface of upper end of cen-
tral nervous system of Crisfatelta. C. General plan of the nervous
system of CtintatelUi. The tentacles are all cut away in one arm
and partly cut off in the other. The position of the alimentary canal
is indicated. D. Side view of a portion of the chief ganglion show-
ing the nerves of the epistome. E. Diagram of sense cells and nerve
bands connected with a single tentacle. F. Diagram of a section
from side to side of the central ganglion showing the cerebral cavity.
A-D, I after Gerwerzhangen from CrisfateUa. E, F, G, and H. Sur-
face longitudinal and cross sections through the ganglion of a fresh
water bryozoan from Oka.
48 Journal (if Entomology and Zoology
From the above review it will be evident that we know much
more about the nervous system of fresh water forms than marine
ectoprocts, and Gerwerzhagen, 1913, has still further extended our
accurate knowledge of the nervous system of fresh water forms.
Most of his information comes from the study of total preparations.
The general form of the nervous system is shown in Fig. 19A.
The cerebral ganglion is connected with the two large ganglionic
cords which have branches to the tentacles by way of the radial
nerves, each of which has two branches. In the upper part of the
(igure is the oral nerve ring while below is the narrower epistomial
nerve ring.
Fig. lOR. shows more detail in the region of the oral nerve
ring and oesophageal plexus. It shows three bands of commissural
fibers running across the cerebral ganglion.
Fig. 19C. shows the general outline of the whole animal with
the tentacles partly cut away. Besides the general nerves there
is the nerve plexus of the base which connects with that of other
members of the colony.
Fig. 19D. is a side view of part of cerebral ganglion. The
nerve supply to the epistome shows on the left.
Fig. 191. shows the nerve supply to the base of a tentacle; two
chief branches enter each tentacle, with sensory nerve cells.
Fig. 19D. shows a diagram of a cross section through the cen-
ter of the cerebral ganglion.
In general then the nervous system of CristeUa may be sum-
marized as follows:
1. The ganglion is hollow with an extension into the two large
ganglion cords.
2. There are two main branches running down each tentacle
one from each adjoining radial nerve from the ganglinic cord. There
are also strands from the bipolar sense cells in the epithelium of
the tentacles. These afferent fibres join the radial nerves on each
side.
;;. There are two nerve rings, the epistomal or dorsal smaller
one and the oral or ventral larger one, each with numerous second-
ary branches.
4. The sense cells in the tentacles, especially are bipolar.
Multipolar cells are also found in the nervous .system and nerve
plexus.
5. There is a ganglion cell network in the wall which con-
nects one member of the colony with another. This network joins
Fig-- 1^". Bryozoa All but, B and C from endoproctans. A. Diagram of the
nervous system and sense cells of Loxosoma. Harmer. B. Longi-
tudinal section of an estoproctan bryozoan from Delage and Her-
ouard. The position of the ganglion is shown by a black area. o.
An avicularian from Biigida showing ganglion after Ladewig. I).
Pedio'llhia showing location of ganglion. E. Ganglion of Pedicellina.
Nitsche. F. Diagram of sense cells in surface of tentacle of Pedi-
cellina. Retzius.
50 journal of Entomology and Zoology
with the similar multipolar network over the surface of the indi-
vidual members of the colony. In the connecting portion of the
colonial wall are no sensory cells so these nerve cells must have a
motor function.
6. The sympathetic system is represented by fine nerves from
the aboral surface of the ganglion to the dorsal and dorso-external
wall of the oesophagus. Ventral fibers also join with the oral nerve
ring by anastomoses.
There is a nerve network over the surface of the alimentary
canal. At the beginning of the oesophagus and extending to the
stomach there is a network of cells and fibers forming a sort of
nerve ring. Further down all parts of the alimentary canal have
a nerve plexus. The nerve net is especially abundant about the
rectum. The function of the sympathetic system seems to be
motor. The sympathetic system in the digestive canal consists of
a nerve network of ganglion cells as well as stands of nerve fibers.
Endoprocta.
Van Beneden, 1845, although he considers PedicelliHa, gives
little or nothing on the nervous system. Kowalewsky, 1867, dis-
cusses the development and Uljanin, 1869, gives the position of
the ganglion in the same genus. Nitsche, 1875, shows the general
position and chief branches of FcdicrUimi. Salensky. 1877, gives
the general location of the ganglion in Loxosoma.
Harmer, 1885, gives one of the best early accounts of the nerv-
ous system of Lnxosoma. He describes a dumb-bell-shaped gan-
glion, bipolar cells on the surface and a median fibrous part.
Nerves pass from the ganglion to the tentacle prominences. There
are many sen.se cells in the tentacles. Silver nitrate was used
to determine the position of the sense cells. The ganglion is devel-
oped from the ectodermic floor of the vestibule and is connected
with a well developed system of peripheral nerves ending in sense
cells bearing tactile hairs on various parts of the body. The adult
has no supraoesophageal ganglion. The nervous system of Lox-
iisoma develops by ectodermic invaginations: the connection be-
tween the two parts is established secondarily.
Foettinger, 1887, represents the nervous system of PcdiciUiiia
l)y a brain more or less comi)letely divided into two lateral lobes.
It is formed by a mass of ganglion cells surrounding a fibrous
center. From the ganglion several pairs of nerves pass.
Seeliger, 1890, gives the development and position of the
nervous system in endoprocts.
Daveni)ort, 189.''>, shows the position of the ganglion in {'»«-
tella.
Nickerson, 1901, in L. ilavcnpoi-ti describes the brain as just
in front of the intestine and above the stomach, between it and the
Pom-jiia Collesjie, ClarcnKint. California 51
floor of the diaphragm. It is elongated transversely, the two
rounded ends being composed of a surface layer of cells with deeper
fibers. Some of the fibers form a commissure. From each end of
the brain two bundles are given off; one on each side passes to the
lophophore. Sonsory bristles were seen from the tentacles. Dorsal
sense organs as described in other forms are absent in this.
Stiasny, 1905, shows the ganglion of PediceUina but with no
detail. Retzius, 1905, shows the sensory nerves in the surface of
PediceUina. These sensory cells bear bristles and are connected
with nerve strands which form a wide network of fibers. Sensory
cells were found in the tentacles.
Assheton, 1912, found the nervous system in two species of
Loxosoma. The branches are figured and sense cells are mentioned
on the hypostome, lophophore and tentacles.
I have been able to study the reactions of two Pacific coast
species of endoproctans. In Barentsia gracilis Hincks, the condi-
tions are much as in PediceUina. The ganglion is small and in the
usual position. The animals are colonial with narrow strands con-
necting the individual members of the colony ; the muscular bases
of each individual cause them to rotate in an active manner. Gen-
eral conditions in Myosoma spinosa Robertson are similar except
that the whole stem is flexible. In Barentsia the polype at the
end of the stem is movable at its stalk. The ganglion is much as
Nitsche describes. There is some indication of sense cells as shown
by Harmer as demonstrated by the methylene blue method although
I never obtained a perfect picture. The tip of the stem is slightly
smaller where it joins the body of the individual and methylene blue
shows bipolar cells at this point. Along the stem there are sensory
pits which are the only breaks in the strong chitin-like covering of
the ten elongated cells of the stem. In Myosoma, in place of the
pits on the skin there are well developed hollow hairs much like
those of arthropods.
Tactile or other stimuli may cause a rotation of the stems with-
out a contraction of the tentacles, but severe stimuli will also cause
the tentacles to contract. Stems with their tips cut from the body
continue to rotate when stimulated. Movements of the body
of the polype on the stems may be caused by tactile stimuli. The
eff"ects of stimulation may be carried from one polype to another
through the connecting stems. One polype in line with others
may be fatigued so that it will not carry the stimuli to others.
The stems and bases of both species seem capable of exciting
movements of the individual as a whole better than the tentacles or
body. In the rotating movements the tentacles are not often re-
tracted unless the stimulus is very severe or the tentacles them-
selves are touched.
The control of movements of the tentacles and body are prob-
ably centered in the ganglion. The excitation to the rotation of
52 jiuirnal of Fntom!)l<)K\ :in»i Zoolog\
the stems is effective through the stems themselves and the pres-
ence of the ganglion is not necessary for these characteristic move-
ments. The conduction from one member of the colony to another
seems more evident than from the base or stem to the tentacle re-
gion, and vice versa.
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VOLUME FOURTEEN _Z_ NUMBER THREE
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
SEPTEMBER, 1922
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT 0/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
The Skui.l of Notothai.omus Torosus — .SVvrr//i Manmon 55
A New Aphis on California Sage — £. O. Esiuj 61
The Nervous System and Sense Organs, X — //'. A. Hiltnn 65
Botered Claremont, Cal.. Post-Office Oct. 1, 1010. as second-class oiatter. under Act of Congress of
March 9. 1870
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The Skull of Notothalamus Torosus
Sarah Marimon
There are twenty-eight bones in the entire skull of Notothal-
amus torosHS. There are, however, only fifteen different kinds,
since thirteen are paired.
These paired bones are : nasals, ectethnoids, maxillaries, fron-
tals, parietals, squamosals, quadrates, ptergoids, occipitals, squamo-
palatines, sphenethnoids, and (of the mandible) dentaries and artic-
ulaires.
The unpaired bones are: the parasphenoid and the premax-
illary.
The premaxillary ( 1 ) is the bone which forms the external
division between the two nares. It consists of a rather thin
dividing bone, which broadens out to form a broad flat base. Dor-
sally the dividing bone diverges posteriorally to form two slender
processes which join at the ends with the premaxillary processes of
the frontal bones, and articulate on the exterior sides with the two
nasal bones. Ventrally the dividing bone broadens out suddenly
into a broad flat base which forms the most forward portion of the
roof of the mouth, and serves as a connection between the two
maxillaries.
The nasals (2) are two irregularly shaped bones, each of which
articulates on the interior side with a premaxillary process, on the
exterior side of the ectethnoid and the maxillary. Anteriorally
the nasals bound the dorsal side of the nasal cavity.
The ectethnoids (3) are two triangular bones located on either
y^l -^
FIGURES
In lettering- these figures I have used the following method:
Each bone on the skull is marked with a large number. Throughout the
figures, each bone goes by its number. When an articulation with a certain
bone is indicated a small figure is used. The bone itself bears a large figure.
When the bone borders on a cavity, or has a free edge, that portion of the
bone is not numbered.
Upper or outer surfaces are indicated by numbers, under or inner surfaces
are indicated by the same numbers prime (1).
5b
Journal of KinonioloK> and Zoology
side of the anterior portion of the skull. Each is so placed that one
half of its surface articulates with the dorsal surface of the skull,
while the other half forms a portion of the side of the skull.
Through the center of the ectethnoid there is a dividing ridge which
separates its dorsal surface from its lateral surface. The dorsal
surface articulates anteriorally with the maxillary and the nasal,
and posteriorally with the frontal. The lateral surface articulates
anteriorally with the maxillary and posteriorally with the frontal ;
a small portion remains between the.se two articulations and this
l^ortion bounds part of the opening into the olfactory fossa.
The maxillaries (4) are long slender bones, which, together
with the premaxillaries bear the teeth of the upper iaw. The
maxillary articulates only anteriorally and diverges posteriorally to
form a long rather slender process extending al)out one-half the
length of the skull. Before the maxillary articulates with the main
body of the skull it diverges on the inner side into two portions;
the upper portion articulates with the dorsal l)ones of the skull,
while the lower portion articulates with the ventral bones. The
hollow depression resulting from these two divergences forms the
opening into the olfactory fossa. Dorsally the maxillary articu-
lates with the nasal and the ectethnoid; and ventrally with the pre-
maxillary and the squamo-iialatine.
The frontals (,5) form a little less than one-half of the dorsal
surface of the skull. They lie in contact with one another for about
two-thirds of their length, diverging anteriorally to form a pair of
short prcmaxillary proce.sses, and diverging itosteriorally 1o form
two processes, — a blunt rather broad parietal i)rocess and a long
slender i)rocess which articulates with the sijuamosal bone. On the
ventral surface of each frontal is an out curving ridge which serves
for the attachment with the sphenethnoid.
PoiiiniKi Ccille"c, Clurcmoiit, Caliti
57
The parietals (6) lie in contact with one another for their
entire length. They are smaller than the frontals, however, they
form less of the dorsal surface of the skull than their real size
would warrant, since at their articulations with the frontals these
bones extend down over them. Posteriorally each parietal articu-
lates with the occipital, and posterio-laterally with the occipital and
the sphenethnoid.
The occipitals (10) are the most posteriorally placed bones in
the skull. The dorsal surface of each is more or less regular, di-
verging toward the median line to form a short rather slender
process which articulates with the other occipital. Anteriorally
the occipital connects with the parietal while posteriorally on the ex-
ternal side, it articulates with the squamosal. The ventral surface
of the occipital is very irregular and the ai'ticulations with other
bones are not continuous. Posteriorally on the external side it
diverges to form a short process, which unites with the squamosal
and forms the posterior corner of the .skull. On the inner aspect
of the ventral surface, adjacent to the squamosal articulation, is a
projecting knob-like process with which the ptergoid articulates.
The most posterior portion of the occipital toward the median line,
diverges to form a knob-like condyle which articulates with the
first vertebra. Anteriorally the occipital articulates with the par-
asphenoid and anterio-laterally diverges to form a projection which
articulates with the sphenethnoid.
The squamosal (9) is a peculiar T-shaped bone, standing in an
almost vertical position in the skull. The bar of the T, forms a
part of the dorsal surface of the skull and articulates at the anterior
end with the frontal bone ; at the posterior end with and up past the
point where the stem of the T diverges, it articulates with the occip-
ital bone. The stem of the T projects ventrally almost at a right
angle, and articulates with the ptergoid and the quadrate.
58
it Einiim()li)j;y and Zoology
A true squamosal bone is sometimes considered not to exist
among Amphibia, and the so-called squamosal bone is considered
to be rather an investing bone on the surface of the quadrate, and
for this reason is sometimes called the jiaraquadrate.
The quadrate (8) is an irregularly shaped little bone with
somewhat the appt-araiice when in position, of a wedge between the
ptergoid and the squamosal. Functionally it serves as a piece in-
terposed between the skull and the mandible, and forming an
articular surface for the latter. The knob-like anterior ventral
end of the quadrate consists of an articular process, fitted with
a socket to receive the rounded knob ( articulare) of the mandilile.
The ptergoid (7) is a spade-shaped bone which projects down-
ward from the ventral side of the skull. It articulates with the
main body of the skull by means of a hollow, rounded process which
articulates down over a knoij-iike projection on the occijiital bone.
Aside from the articulation with the occipital, the ptergoid articu-
lates posteriorally with the (luadrate and the squamosal.
The squamo-palatines (1:1) are long rather slender bones, flat-
tened anteriorally. At about one-third of their length, from the
anterior end, they articulate dorsally with the parasphenoid and
project down onto that bone for the remainder of their length.
These projections are provided with teeth along the median line.
Anteriorally the squamo-palatines articulate with the premaxillary
and the maxillaries.
The parasphenoid (11) is the flattest and most extensive bone
in the skull, and forms nearly the whole floor of the brain case, and
at the same time the roof of the mouth. It is nearly the shape of
a parallelogram with rounded corners, but it is a little broader in
the optic region and becomes somewhat narrowed anteriorally. It
Pom )na C()llfL;e, Clnrcniont, California 59
has no especial markings or features other than the impressions
made by the bones which come in contact with it. On its ventral
surface are two long narrow impressions left by the squamo-
palatines.
The sphenethnoids (12) are the bones which serve as walls to
hold apart the dorsal and ventral surfaces of the skull. They are
rather long bones, about three-fourths as long as the parasphenoid.
They articulate posteriorally with the occipitaLs, their dorsal edge
articulates with the frontal? and parietals, their ventral edge with
the parasphenoid. Anteriorally they bound a portion of the open-
ings into the occipital fossae.
The mandibles of Nofofbakniius forosif>i are each composed of
two bones, the dentary and the articulare.
The dentary (14) is that part of the mandible which bears the
teeth. It is a long slender, curved bone, articulating anteriorally
with the other dentary, and widening out posteriorally to articulate
with the articulare.
The articulare (15) is that part of the mandible which diverges
posteriorally to form a rounded knob which fits into the articular
socket of the quadrate. Anteriorally, on the median side it fits
down into the dentarv bone.
A New Aphis on California Sage
APHIS HILTONI n. sp.
(Figure 1)
By E. 0. Essig, Division of Entomology
University of California
Apterous Viviparous Female. —
(Figure 1, A). Length 1.3 mm., width of abdomen 0.9 mm.
Prevailing color pale green, the dorsum partially covered with a
fine white powdery wax which is arranged in minute pore-like or
mosaic rings. The areas not so covered appear dark in the illus-
tration. There are numerous black pigmentations dorsally and lat-
erally on the epidermis of the mounted specimens. The cornicles,
Cauda and anal plate ; all of the legs excepting the basal three-
fourths of the tibiae ; and antennal articles, VI, V, II, I and the tip
of IV are black or dusky. The remainder of the antennae and tibiae
are yellow. The rostrum extends slightly beyond the base of the
abdomen. The antennae are shorter than the body, the relative
lengths of the articles being :
I. 0.065 mm., II. 0.055 mm.. III. 0.227 mm., IV. 0.167
mm., V. 0.155 mm., VI. 0.280 mm., (base 0.130 mm., spur 0.150
mm.), total length 0.949 mm. There are the usual sensoria on
articles V. and VI. The prothoracic tubercle is well pronounced.
There is also a well defined pair of antei'ior and a pair of posterior
abdominal tubercles (Figure 1, A. tub. i, ii, iii). The tarsi are
small and one-third as long as the cornicles. (Figure 1, At.).
The cornicles are black, cylindrical and somewhat tapered towards
the tip, straight, slightly imbricated; 0.37 mm. long, and 0.06 mm.
wide at the base. The cauda and anal plate are black (Figure 1,
A. Cauda).
Winged Viviparous Female. —
Length 1.20 mm., width of abdomen 0.56 mm. Prevailing
color black with abdomen and legs dusky yellow. The dorsum may
also be partially covered with a fine white powdery wax. The
antennae (Figure l.W. ant) are dusky to black throughout, the
length of the different articles : I. 0.070 mm., II. 0.050 mm.. III.
0.200 mm., IV. 0.155 mm., V. 0.153 mm., VI. 0.280 mm. (base 0.125
mm., spur 0.155 mm.), total length 0.908 mm. Article III usually
has four or five large circular sensoria along the lower side, but
there are sometimes six. The usual sensoria occur on V and VI.
The rostrum reaches to the second abdominal segment. The pro-
thoracic and abdominal tubercles are much like those in the apter-
ous form and are illustrated in Figure 1, W. tub. The wings (Fig-
ure 1, W.) are normal in venation as illustrated. The lengths are:
62 Journal ot Rnti)m<ili)t;v and Zoology
primary 2 mm., secondary 1.2 mm. The cornicles are black, imhi-
cated, cylindrical, somewhat larger near the base, the outer margin
straight, the inner margin as illustrated (Figure 1, W. corn. ) . The
length 0.10 mm., greatest width 0.05 mm. The cauda and anal
l)late are black and as illustrated (Figure 1, W. cauda).
Relationship — This species has been carefully checked with
Aphis reticulata Wilson, A. oregonetwiis Wilson, A. hermistonii Wil-
.son, A. tridentatac Wilson, A frigidae Oestlund, and Aphis artc-
m;.sToto Williams occurring in Oregon on Arteviisia tridcntata, and
does not agree with any of them or other closely related species.
Host — The species occurs in dense colonies on the apical twigs
of old man or California sage, Artemisia calif ornica Less.
Locality — In Laguna Canyon one-half mile above Laguna
Beach, California.
Date of Collection — July 13, 1921.
COTYPES — The above description was made from a series of
cotypes consisting of ten slides and over one hundred mounted indi-
viduals. The cotypes are in the author's collection.
The species is named after Dr. Wm. A. Hilton, Professor of
Zoology, Pomona College, under whose su])ervision. inspiration and
energy a most wonderful type of biological in.struction is being
given each summer at the Laguna Beach Laboratory.
Figure 1. — Aphis hiltoni n. sp.
A. Apterous viviparous female; A. tub., body tubercles of
apterous female; i, prothoracic; ii, front abdominal; iii, posterior
abdominal; A. corn., apterous cornicle; A., cauda, apterous cauda
and anal plate; At. t., apterous tarsus: A. ant., apterous antenna;
W, wings; W. ant., antenna of winged female; W. corn., cornicle of
winged female; W. cauda, cauda and anal plate of winged female;
W. tub., body tubercles of winged female; i. prothoracic; ii, front
abdominal; iii, posterior abdominal.
X. Phoronida and Actinotrochia
Wright in 1856 described the first species of Phoronis. Dyster,
1858, suggests two oesophageal gangha. He found that the crea-
tures were not particularly sensitive to light. I have a reference
to a paper by Kowalevsky, 1861, on the anatomy and development
of Phoronis, but as I have not seen it or a review of it I do not know
how much the nervous system is considered.
Caldwell's publication of 1883 is the next paper of importance.
He describes the nerve processes in connection with the ectoderm;
both fibers and ganglion cells occur in the ectoderm. There are
concentrations of the nervous tissue about the mouth to form a
post-oral nerve ring; the anus is outside of this. The ring forms
a line along the base of the tentacles formed like a horse shoe. In
front of the ring is a pair of sense organs, the ciliated pits in the
concavity of the lophophore on either side of the anus. There is an
epithelium here with sense cells, ganglion cells and nerve fibers.
The nervous system is further continued on the left side from the
dorsal part as a cord or strand just outside the basement membrane.
Mcintosh, 1888, in P. buskii describes a similar epidermal sys-
tem concentrated about the mouth to form a post-oral nerve ring
with the anus outside. The ring follows the line along the base of
the tentacles and to the "ciliated pits" or concavity of the lophophore
on either side of the anus. The nervous system has sense cells
and ganglion cells and nerve fibers. On the left side is a cord
through the body. The left longitudinal nerve tube or tubes of
Caldwell is not described.
Andrews, 1890, in a new species describes the "glandular pit"
of the lopophore and a large "nerve rod" on the left side, solid and
surroundecl by epidermal cells. It seems to have a fibrillated or
possibly only a coagulated structure. The rod extends through a
considerable distance and ends in a peculiar ring of epidermal nerve
substance about the mouth. At this region there are two symmet-
rically placed nerve rods but the right is short.
Benham, 1889, finds the nervous system immediately below the
epidermis as Caldwell was first to observe. Passing aborally from
the lopophore ridge the basement membrane is seen to separate
from the epidermis by a narrow ground substance not readily
stained. In this granular substance are a few rounded nuclei be-
longing to small nerve cells. Fibers are also found coming from
the epithelial cells of the surface. This nerve band follows the
ridg« of the lophophore rassing around on the oral side and curves
at the side of the nephridial ridges following the spiral course of the
lophophore. It always keeps along the outer edge of the tentacles.
From this band nerve tissue goes to each tentacle passing along its
inner surface. A nerve goes to each nephridium and a nerve layer
to the epistome, this being the only dorsal part of the nervous
ftf) Journal of Entomologi,- and Zoology
system. There are no concentrations any place to form a ganglion ;
tilt' chief nervous system lies vent rally. Two longitudinal nerve
tubes or nerve hands are described running the length ot the body.
The nerve strands may give the appearance of tubes due to shrink-
age; epithelial cells seem to compose it and the tissue does not look
like nervous tissue. These longitudinal tubes or nerves may be
some sort of sense organ.
Cori, 1890, adds nothing to the general knowledge of the nerv-
ous system.
Torrey, lUll, in P. iHirilica gives a partial description of the
nervous system as like that of P. architecta with the exception that
"The two longitudinal cords which are of e.xceedingly unequal
length, instead of continuing in the nerve ring of the lopophore, are
continuous across the median line at the level of the median mass
of ganglion cells. The loop thus formed is closely applied to the
latter and touches the lophophore nerve on each side of the rectum,
apparently without fusing at either point." I have found no such
condition in several good series of well stained Ph(ir<))ns ixicitiva.
Either this was an individual difference or Torrey's material was
poorly fixed.
Schultz, 1903, discusses the regeneration of the central nervous
system.
De Selys-Longchamps, 1907, described the circular nerve ring
and ganglion and the lateral nerve of Caldwell on the left.
Pixel], 1912, discusses two new species of Phoronida.
In Phoronis vanconverensis, there is the usual ring of nervous
tissue at the l)ase of the lophophore; from it five nerves continue up
the tentacles. Across the dorsal surface in front of the anus is a
large ganglionic mass composed of fibers and cells with large nuclei.
This tissue is everywhere in intimate relation with the inner ends
of the epithelial cells. In some sections two small lateral nerve
cords ran along the right and left sides of the body close to the
l)oint of attachment of the lateral me.senteries and projected into
the basement membrane. He describes these as, "punctated tis-
sue." They are very short. Nervous tissue was found in the
center of the pit at the proximal end of the body and also along the
/ilimentary canal on the outer side of the epithelium especially
marked in the region of the oesophagus opposite the nerve ring.
Gilchrist, so he says, suggests this patch as an organ of taste.
Phoronapsiii hamesi has a similar condition of the nerve ring
luit the ring is narrower and more elongated than in PliorouiK. A
conspicuous nerve cord extends down the left side. In the neph-
ridial region it is sejiarated from the ei)ith('lium and embedded in
the basement membrane; after i)assing inti'rnally to the nei)hridial
duct it turns outwards and rejoins the epithelium a little to the
oral side of the lateral mesentery. From here it extends as a con-
Ponicina Ccille^c, Clait-ninnt. Cnlifciinia
67
spicuous cord in contact with the epithelium and projecting slightly
into the basement membrane. The center is of a clear substance
and about this center are nerve cells.
Harmer, 1917, in Phoronis oralis gives the position of the
nerve ring which he shows thickened on the dorsal side.
I have had some opportunity to study P. pacifica and a species
of Phoronapsis. It is quite important in studying the serial sec-
tions of this group that rather perfect preparations be available, a
condition not altogether easy, as sand often interferes with gootl
sections. However a number of perfect preparations were ob-
tained.
In general I found the nervous system much as already indi-
cated by the many of the others. In P. pacifica I found central
nervous svstem to have its chief concentration a little below the
Fig. 21. A. Section showing position of nervous system off /ioi Jin's- after
Schneider. B. Nervous system in Phoronis after Schneider. C.
Diagram of a reconstruction of the nervous system of Phoi-oiiif! show-
ing longitudinal nerve cord on the right. Not all of the nervous sys-
tem going to the tentacles is shown at the left. D. Section of nerve
cord with epithelial cells on the outer surface and basement mem-
brane in dark below. E. Actinotrocha larva showing the nervous
system after Inedia. F. Diplochorda, after Masterman.
68 Jdurnal dI^ Ent()m()lot;\ ami Zoology
level of the anal opening and the nephridial tubules. This thickest
portion of the nervous system directly continues with the epithelium
•of the surface of the body and is dorsal to the anal papilla in the
depression caused by the anal prominence; from here the thicken-
ing passes toward the tentacles sending fibers to the jophophore and
the tentacles. The lophophore depression on each side marks off
the chief thickening of the nervous system. This central part,
although continuous with the epithelium is made up of a distinct
mass of fibers and cells. At this point three chief centers of cells
are found among the fibers while out laterally strands run to the
lophophore dcfressions and out to the tentacles. There is quite a
mass of fibers and cells in the region of the lojjhophoral depression.
Running out vertrally on the left side just medial to the lophophoral
depre.ssion and between it and the left nephridium is the clear cord
of unknown function noticed first by Caldwell. This cord sur-
rounded partly by cells comes to run farther ventrally until it
passes through the basement membrane of the liody-wall and comes
to lie .iust under the ejiithelium. This end does not seem to be of
nervous tissue, although it is connected with the central part of the
nervous system.
If I understand Torrey's descriiition aright his material mu.^^t
have been too poorly fixed to show the relationship of the nervous
system for in well preserved sjiecimcns the cerebral nervous sys-
tem is continuous laterally with the lophophoral organs as well as
with any lateral or longitudinal extensions of the nervous system.
My observations both on Pharonis and Phoroiiapt^is agree closely
with tho.se of Pixell. In Phoronapsis the centra] nervous system
seems more elongated, as Pixell found.
With the exception of the central part of the nervous .system
the nerve cells are not clearly different from the eiiithelial cells, but
careful study shows at the liases of the cells as well as farther down,
nerve cells with their fibers directed into the basal mass of fibers.
Ill the epithelium are bipolar cells, some of which may be sen.sory,
although many of the ]n-ominent strands are tho.se of supportive
cells.
ACTINOTROCHA.
It seems best to consider the larval stage of Plii)r())iis iiriefly
at this place. Schn(>i(ler, 18(i2, in his discussion of the develop-
ment of Arti)Hitri>rh(i does not consider the nervous system. Cald-
well has the first work of imiiortance but his account, according to
MacBride, implies that the apical jilate and adjacent ganglion of
the larva are lost, and the cerebral ganglion of the adult mu.st be a
new structure. But in every trochophore so far studied the apical
plate with its ganglion forms the material which persi.sts to the
adult condition.
Pomona College, Clareniont, California 69
Masterman's paper of 1898 is a very important one. He men-
tions Wagner, '47, as the first to describe the nervous system. Mas-
terman describes a central ventricular ganglion in the mid-dorsal
line at the base of the prae-oral lobe, composed of ganglion cells
and fibers. The ganglion is a proliferation of the inner cells of the
epiblast. Nerve tracts radiate in almost every direction.
The nervous system may be summarized as follows :
1. Central ganglion in front collar region and between this
and the prae-oral lobe. The epiblast in front is depressed to form a
neuropore.
2. A ring about the posterior part of the collar is continued
dorsally and ventrally giving olT fine double groups of nerve tracts
to the anal end of the body.
3. Groups of fine nerve tracts continued dorsally along the
trunk from the anterior end of the collar.
Fig. 22. A. Section through body and central nervous system of Plioronis.
B. Small portion of lopophore showing depression. C. Small por-
tion of the nervous system of Phoronis enlarged to show nerve cells.
70 jiiunial iif Entiimolojiy ami Zoolopy
4. A ring about the anal end of the trunk into which dorsal
and ventral tracts lead.
5. A ring about edge of prae-oral lobe, joined at each side to
the ganglion and in median front region by three main tracts run-
ning in mid-dorsal line.
6. A diffuse plexus of fibers at the base of all the epiblastic
layer, including fibers of ventral collar region, which pass forward
and dorsally to meet the ganglion.
Ineda, 1901, found no collar, nerve ring or dorsal or ventral
commissure in the larva. He also failed to make out presence of
(he peri-anal ring. If present it is represented by a small number
of parallel fillers. The main nerves were three in number close to
each other and parallel along the mid-dorsal line of the trunk but
confined to only a few sections posterior to the first pair of tentacles.
There was found however a very complex and beautiful system of
nerve fibers seen on the prae-oral lobe. Fibers are very numerous
and fine and radiate from the ganglion on all sides towards the free
margin of the prae-oral lobe. In the median line and anterior to
the ganglion fibers are three long parallel strands on which the
apical sensory spot is situated, not far from the ganglion. After
passing through the sensory spot .strands fray out into fine fibers
which continue to the free margin of the prae-oral lobe. Fibers
from (he ganglion do not show a regular radial arrangement, but
ari.se from the lateral edge of the ganglion and soon take an anterior
direction. Sometimes near the ganglion there is an anastomosis of
fibers, but probably more apparent than real. There are nerve end-
ings in the prae-oral ciliated belt. There is probably an incomplete
development of nerve elements in the collar and trunk region. He
finds no neuropore and believes that Ma.sterman's structure is due
to contraction.
De Selys-Longchamps, 1902, gives a rather complete descrip-
tion of the nervous system. The central ganglion is a dorsal ex-
pansion of the epidermis with fibrillar substance below the surface.
The depression which Mastcrnian calls neuropore is not such a
structure. There are three cords of the nervous system, the median
is most developed. The apical organs are organs of sense.
BIBLIOGRAPHY
.Andrews, E.
1900. On a New American species of the remarkal)lc animal I'horonis.
Ann. Map. Nat. Hist. vol. 5, pp. 44.'>-l49.
Benham, W. B.
188il. The Anatomy of Phoronis australis. Q. Jour. inic. so. vol. :!(>. pp.
125-158, pi. 10-1.-?; N. syst. pp. I.'JS-ISS.
Caldwell. W. H.
18S:i. Preliminary note on the structure, development and affinities of
Phoronis. Proc. Roy. soc. vol. 34, pp. 371-.383, 1 fig.
Pomona C(j1 lege, Claremoiit, California 71
Cori. C. J.
1890. Unterauchungen uber die Anatomie und Histologic der Gattung
Phoronis. Zeit. f. wiss. Zoo), vol. 49, pp. 28n-.';68, pi. 22-28.
Dyster, F. D.
1858. Notes on Phoronis hippocrepia. Trans. Linn. soc. vol. 22, pp.
251-255.
Goodrich, E. S.
1903. On the body cavaties and . Nephridia of the Actinotrocha larva.
Q. Jour. mic. soc. vol. 47, pp. lO.S-121, pi. 8-9.
Harmer, S. F.
1917. On Phoronis ovalis Strethell. Q. .Jour. mic. soc. vol. 62, pp. 115-
148, pi. 7-9.
Haswell, W. A.
1882. Preliminary note on Australian species of Phoronis (P. australis).
Proc. Linn. soc. N. S. Wales, vol. 7.
Ineda, J.
Observations on the development, structure and metamorphosis of Actino-
trocha. Jour. Coll. sc. Imp. univ. Tokyo, vol. 13, pp. 507-592, pi.
25-30.
Kowalevsky.
1867. Anatomic und Entwicklung von Phoronis. St. Petersburg.
Mcintosh, W. C.
1881. Notes on Phoronis dredged by H. M. S. Challenger. Proc. Roy.
Soc. Edinb. vol. 11, pp. 211-217.
1888. Report on Phoronis buskii n. sp. dredged during the voyage of
H. M. S. Challenger. Zool. vol. 27. part. 7.''>, pp. 1-27, pi. l-.l.
Masterman, A. T.
1897. On the Diplochorda.
1. Structure of Actinotrocha. 2. Cephalodiscus. Q. Jour. mic.
sc. vol. 40, pp. 281-366, pi. 18-26.
1300. On the Diplochorda. Ill The early development and anatomy of
Phoronis buskii. Med. Q. Jour. mic. sc. vol. 43, pp. 375-418, pi.
18-21.
1901. Prof. Roule, upon Phoronidea. Zool. anz. Bd. 24, pp. 228-233.
Pixell, H. L. M.
1912. Two new species of Phoronidea from Vancouver Island. Q. Jour,
mic. sc. N. S. vol. 58, pp. 257-284, 16 text figs.
Roule, M. L.
1897. Sur le development des feuillets blastodermigues chez les Gephy-
riens tubicoles (Phoronis sabbatieri n. sp.) C. Ac. d. sc. Paris
vol. 110, pp. 1147-1149.
1900. Remarques sur un travail recent de M. Masterman concernment le
developpement embryonnaire des Phoronidiens. Zool. anz. vol.
23, pp. 425-27.
72 Journal ot Fintoniolopi' and Zoolnp'
(le Selys-Lonjrchamps, M.
1902. Recherches sur le development des Phoronis. Arch, de Biol. vol.
18, pp. 495-.197, pi. 22-24.
1907. Phoronis. F. & Flora. Golf. Neap. 30e Monog. pp. 1-280, 1 te.xt
fig. 12 pi. N. syst. pp. 49-61.
Schneider, A.
1862. On the development of Actinotrocha branchiata. .\nn. Map. nat.
Hist. vol. 9, 3 d. .ser. pp. 4f6-7.
Schultz, E.
1903. Aus Gebiete der Regeneration. Zeit. f. wiss. Zool. vol. 7.5, pp.
390-420, pi. 27-28, and pp. 472-494, pi. 33.
Torry, H. B.
1901. On Phoronis Pacifica sp. nov. Biol. Bull. vol. 2, pp. 283-288, figs. 1-5.
Wagner.
1847. Actinotrocha. Arch. f. anat. u. Phys. pp. 202-206.
.^ir NOV 17 1939 ' W!'
VOLUME FOURTEEN NUMBER FOUR
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
DECEMBER, 1922
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT of ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
On the Occurrence of Polvgordius Adult at Laciuxa Beach —
W. A. Hilton 73
Insect Notes from Lacuna Beach, California — E. O. Essig 75
Nervous System and Sense Organs, XI — //'. -7. Hilton 79
Sutered Claremont, Cal.. Post-Offlte Oct. 1. lUlo. as second-class ojatler. under Act of Cuiigrens of
March S. 187V
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The Occurrence of Polygordius Adult
at Laguna Beach
William A. Hilton
For a number of years now we have taken Branchiostoma just
off shore in rather coarse sand, but it was not until the summer of
1920 that we began to look for archiannelids. A few doubtful
specimens were obtained from sea weeds but nothing that we could
be sure were the animals sought. We never thought to search the
sand in which Branchiostoma was taken until after reading in the
monograph on Polygordiw^ how the creatures were obtained near
Naples. With the hint that these animals were sometimes asso-
ciated we examined with great care some hundreds of pounds of
coarse sand in which some few Branchiostoma had been found and
from this two specimens were obtained, one dead and one living.
These were without question of the genus Pnlygordius although to
make the matter more certain sections were made. Although the
genus is certain, the species remains undetermined because the
caudal ends of the animals were not perfect.
The living specimen was very active. At first it was taken to
be a rather long round worm but the characteristic antennae at the
head region caused it to receive more attention.
So far as I can tell, this is the first record of the adult of Poyl-
gordius being found in North America in its natural environment
at least, for some have been reared from the larva! forms at Woods
Hole.
(Coyitrihvtion from the Zologiral Laboratory of Pomona Col-
lege.)
Insect Notes from L^aguna Beach,
California
By E. 0. Essig, Division of Entomology
University of California
The following notes were made during the Summer Session
at the Pomona College Marine Laboratory, Laguna Beach and
vicinity during June and July, 1921.
ORTHOPTERA^
Two earwigs, Anisolabis aniiidipci^ Lucas and A. maritima
Brun., were commonly taken in the canyons in damp places under
stones, logs and in wet leaves. The former occured under stones
close to the creeks.
The cockroach, Arenivaga (HomoUxjamia) erratica (Rehn),
was taken under a large stone. The specimen taken was appar-
ently full grown and a wingless male. A winged female was also
collected.
The ra?L\\i\ds,Stagmoynayitis califoruicn R & H and Litaneidria
obscura Scudd., were both taken on the hills near the ocean beach
during July 1921.
The common tree cricket in the Laguna Beach region proves
to be Oecanthus nigricornis var. argentinus Sauss. A number of
these were taken during July.
The red Jerusalem cricket, Stenopelmattis fuscus Hald., was
taken in a rotten log in Niguel Canyon. The common species at
Laguna Beach which regularly traverses the streets at night and
may often be found in the morning, is S. longispina Brunner
(Syn. S. irregularis Scudd.).
The large blue-winged grasshopper, Leprus glaucipennis
Scudd., proved to be a match for the most active entomologists and
eluded many a net. The species measures from 2 to 214 inches
long and the color matches perfectly the color of the soil on the
hills, back from the ocean where it occurs. The blue under-wings
easily characterizes it.
ITHYSANOPTERA
Western grass thrips, FrcoikUnieUa (Euthrips) occidentalis
(Pergande)". A pale yellowish-brown species was abundant in
the heads of Junciis xiphiodes Meyer growing in fresh water at the
mouths of the canyons near the ocean.
' Determined by A. N Caudell. Bureau of Entomology, U. S. Dept. of Agriculture.
-Determined by A C. Morgan, Bureau of Entomology, U. S. Dept. Agriculture.
76 Journal of Entomology and Zoology
The Christmas berry thrips, Trichothrips ilex Moulton, occurs
ill all stages upon the tree nialva, Malrastrum fascicidatum (Nutt.)-
The young are bright cardinal red with the head, antennae, dorsum
of prothorax, legs, and last abdominal segment black. The adults
are entirely black. The insects feed on the stems and under-
sides of the leaves and the brilliant red nymphs are often present
in considerable numbers.
This species also attacks the Christmas berry, Heteromeles
arbutifoUa (Lindl.) and a variety Trichnthrips ilex dumosa Moulton
occurs in southern and central California on scrub oak, QiiercK.f
dumosa Nutt.
HEMIPTERA
The Crackling cicada, Cacama crepitants (Van Duzee).- One
of the most interesting insects in the hill region is the crackling
cicada, so-called from the various crackling sounds intermingled
in the long sonorous buzzing or droning which is at times so deaf-
ening. When captured they make a terrific high-pitched noise.
The adults may be observed resting near the tops of various shrubs,
but appear to prefer the California sage, Artemisia, ealifdruica
Less.
The black scale, Sainsetia olenc (Bern.), is abundant at Laguna
Beach, having been dispersed far over the hills infesting many
native plants including the California sage, Artemisia califnrnica
Less., willows (Salix spp.), and the lemonade or sour berry, Rhus
i)ite(irif(Aia B. & H.
The Cabbage Bug, - Murgantia histrionica Hahn.- The native
black pha.se of this species, described as M. nicirirati'< by Cockerell,
occurs in great numbers upon the wild mustard. Brassica cam-
pestris Linn., and more particularly upon the wild bladder-pod,
Iso7neris arhorea Nutt.. growing on the sea coast hills and in the
valleys of Southern California. On the latter plant it overwinters
and survives the dry years when the mustard fails to appear. The
writer lielieves that the above form of the cabbage bug has long
been a resident of Southern California where for ages it has sub-
sisted upon the two plants listed and .should be considered as a
native insect.
The eggs are often heavily parasitized by a minute black
encyrtid, Ooencyrtus john-^oiii (Howard) '. Adults of this parasite
were reared from eggs taken chiefly from the wild bladder-pod
growing on the hills near the ocean from Balboa Beach to San Juan
Capistrano. They issued in greatest numbers during the month
of July.
' Detrrminotl l>v A. H r.nhiin. Huienu of Kntomolonv. U. S. Di'pt. uf Auricultur
Pomona College, Claremont, California 77
DIPTERAL
The common kelp fly, FucclUa rufitibia Stem, was particularly
abundant on decaying kelp along the beach during the summer.
In some instances the flies completely cover the masses of seaweed
and rise in clouds when disturbed. It would be interesting to know
the larval habits of this species.
The lemur syrphid, Baccha lemur 0. S., was reared in con-
siderable numbers from Erium licktensioides Ckll. on California
sage, Artemisia californica Less., which was abundant in the
Laguna Beach Canyon.
The small gray leucopis, Leucopis griseola Fall., was reared in
immense numbers from the leaves of muskmelon vines which were
severely infested with the melon aphis, Aphis gossypii Glover. The
small larvae and pupae were abundant on the undersides of the
leaves. That a large proportion of the muskmelon vines growing
along the ocean between Laguna Beach and San Juan Capistrano,
were not entirely destroyed, may be credited to the efficaceous work
of the larvae of this fly. I have never seen a predaceous maggot so
numerous.
LEPIDOPTERA
The Sycamore borer, Synanthedon (Aegeria) mellinipennis
(Bdv.).'- The work of the larvae of this moth on the trunks of
the Western Sycamore or plane tree, Platanus racemosa Nutt., is
very characteristic, consisting of numerous tunnels in the inner
bark and the expulsion of quantities of reddish-brown frass which
collects in the crevices of the bark and around the bases of the
trees, at once calling attention to the presence of the insect.
The infestations occured on large trees and was confined to
the trunks from the ground to a distance of about six feet. Many
of the trees were infested with great numbers of caterpillars, but
no evidence of serious injury to the general health of any of the
infested trees was noticeable. The moths mimic in color, size and
flight the common yellow jacket, Vespa gcrmaiiica Linn. Indeed
so great was the resemblance that the moths hovering about the
tree trunks were first thought to be yellow jackets until they
alighted.
A single grove of western sycamore, comprising some fifty
trees, in Niguel Canyon was the only one observed to be infested by
this moth, although there were numerous other trees in the difl'erent
canyons around Laguna Beach.
The western sycamore is apparently the native host of this
species, which is recorded from California and Colorado, without
previous host records.
' Determined by J M. Aldrich. U. S. National Museum.
■ Determined by Ausust Busck. U S. National Museum.
7S Journal of Kntomoloy) ami Zoology
HYMENOPTERA
The Yellow aiul Black Mud-dauber, Sceliphron serrillii Le-
peletier.-This interesting dauber is common along all of the streams
in the vicinity of Laguna Beach. The elongated mud cells about
one inch long are built singly or placed side by side in series of
from two to four and the whole covered with a continuous layer
of mud completely ol)literating the outlines of the individual cells.
The cells were commonly ijlaced on the undersides of large rocks
or boulders in the near vicinity of the fresh water streams and
often at the mouths of the canyons near the ocean. The nests
were stored chiefly with yellow and brownish-gray crab spiders.
In the cells and attacking the larvae of the mud-daubers was
often found the maggot of a tachina fly, which proved to be
Pacliijaphthnlmua tioridoisix Townsend . The adults of this most
interesting fly escaped from the masonry cells by the expansion
and retraction of an inflatable bladder-like organ in the front of
the head (ptilinum?) which was used to moisten the mud and then
scrape it away. Adults confined in glass vials were easily observed
to continually endeavor to work their way through in this manner.
Not all of the ffies appeared to possess or to use such an organ, but
whether or not this is a sexual characteri.stic was not determined.
The fire ant, SolenopsiN geminata Fab., was perhaps the com-
monest ant in the vicinity of the laboratory. During July the
ants were swarming from their ground nests in great numbers.
The workers are small, entirely reddish or with small rounded black
abdomens, the winged females are reddish throughout while the
winged males are black.
• Determined liy J. M. Aldilch.
XI. Brachiopoda
Perhaps in no group of animals is our knowledge of the general
arrangement of the nervous system in such an unsatisfactory con-
dition. Various published accounts are not altogether in accord
even when the same species is studied.
Owen, 1835, seems to be the first to detect the nervous system.
He describes white filaments which traverse the visceral cavity and
end in muscles.
Huxley, 1854, considers the nervous system to be a ring of
nervous tissue about the oral opening.
Gratiolet, 1857, 1860, describes a considerable mass of gan-
glinic material encircling the oesophagus but reduced to a small
ring on the upper side of the oesophagus.
Hancock, 1859, says that the nervous system is easily seen but
not clearly defined. In one form studied five centers of nervous
tissue were found about the oesophagus, three of which were large
enough to be called chief ganglia. He did not find a pallial nerve
described by Owen.
Van Bemmelen, 1883, has a more detailed account of the nerv-
ous system. According to this author there is a pair of infra-
oesophageal ganglia and two true supra-oesphageal centers. From
both, nerves run to the arms. The nerve centers are composed of
very small ganglion cells and fibers ; the peripheral nerves are com-
posed of straight fibers.
Beyer, 1886, describes a commissural ring surrounding the
oesophagus at its junction with the stomach, in Lingula. There
are nerve centers in the ring as follows : one central, two dorso-
lateral and two ventro-lateral, these last being the largest. All
centers are below the ectoderm and the .nerve cells communicate
with the surface.
Blockmann, 1892-3, gives quite a complete picture of the dis-
tribution of the ganglia and chief branches. In his work the lat-
eral ganglia are widely separated and little emphasis is given to
any supra-oesophageal center.
Delage and Herouard, 1897, give quite an extensive account of
the nervous system. In their general account they speak of a sim-
pler nervous system presuming to some extent embryonic condi-
tions of connection with the epidermis. There is a large peribuccal
collar formed'of two dorsal cerebral ganglia and a ventral ganglion
much larger and a little bilobed, with a pair of fine connectives.
From the cerebral ganglia nerves go to the arms. From the ex-
tremity of the connectives a pair of nerves run to the cirri. Nerves
in the arms anastomose and form a nlexus of fibrous cells just under
the epidermis. The ventral ganglion gives off, at its posterior
angle, a pair of dorsal pallial nerves which run to a corresponding
80
Journal lit Knt(imol()f;y and Zoology
lobe of the mantle. From the anterior angle a ventral pallial nerve
soon branches into two, one for the dorsal lobe of the mantle and
one for the corresponding adductor muscles. It is probable that
these nerves also go to the muscles and viscera. In the ventral re-
gion is a ple.xus formed by the ventral pallial nerves. In the
mantle the pallial nerves form a plexus with ganglion cells.
There are no positive organs of .sense ; there are neither eyes
nor otocysts. Probably the margins serve as organs of touch. The
cirri are probably for tactile sense, possibly olfactory. They have
a rich nerve plexus.
Stomach papillae Joubin, 1886-92, suggested as gustatoi-y, and
the terminal papillae of the mantle Sollas, 1887, believed had a
tactile function.
In Ecardia, Delage and Herouard give a separate account. A
single pair of ganglia are situated very low and at the external
14-1^ ^}-
P'ifc. 2.T. Nervous System of Brachiopoda. A. Diagram of the nervous sys-
tem from the ventral side showing the ganglion and chief nerves
after Blochmann. Much modified. B. Diagram of the nervous sys-
tem of n brachiopod, after Brammelon. C. Position of the nervous
system shown in position. Diagrammatic. D. Diagram of LinguUt
showing ganglia in dark. E. General plan of the nervous system.
F. Plan of the central nervous system. G. Nerve plexus.
Pomona College. Claremniit. California 81
border of the superior adductor muscles. A large ventral commi.si-
sure unites the ganglia under the oesophagus. Each ganglion fur-
nishes the following nerves: (1) to the adductor inferior muscle,
a nerve with a little branch to the internal oblique muscle, (2) a
nerve to the dorsal part of the mantle. (3) a nerve to the ventral
part of the mantle, (4) a nerve to the arm, (5) branches which join
with the ventral oesophageal commissure, (6) several nerves form-
ing the dorsal nerve commissures. The dorsal commissure has
nerves going to the cirri.
All nerves are under the skin. Cirri are probably organs of
touch.
Heath, 1889, has found sensitive striae formed by high epith-
elial cells connected with the ganglion cells. These areas are along
the middle line on the ventral side.
In spite of fragmentary and conflicting evidence the following
seems clear as the nervous system of brachiopods :
A nerve ring surrounds the oesophagus; this is enlarged on
the dorsal side in a small inconspicuous ganglion near the base of
the lip. A larger suboesophageal ganglion is the thickening on
ventral side. The ventral ganglion and perhaps the dorsal retain
their primitive connections with the surface layer of the skin. Both
ganglia give off a nerve each side to the arms and along the base
of the tentacles and lips. The ventral ganglion also gives off nerves
which supply the dorsal and ventral folds of the mantle and the
muscles. In some cases the dorsal ganglion seems to be repi'e-
sented by a dorsal band only.
Sense organs are doubtful: the margins of mantle and cirri
may have a tactile function and the epithelium on the surface of
the ganglia have been suggested as olfactory areas.
BIBLIOGRAPHY
Bemmelen, J. G. Van
1883. Untersuchungen ueber den Anatomischen unci histologishen Bau
der Braehiopoda Testicardinia. Jen. Zeit. f. Nat. Bd. 10, pp. 88-
161, pi. 5-6.
Beyer, H. G.
1866. A Study of the Structure of Lingula (Glottidia) pyraniidata
Stein. J.'H. Univ. Stud. vol. 3, pp. 227-265, pi. 14-17.
Blockmann, F.
1892. Ueber die Anatomie und die verwandtschaftlichen Beziehung der
Brachiopoden. Arch. d. Fren. d. Nar. in Meek. Rest. pp. 37-50.
1893. Anatomische Untersuchung ueber Brachiopoden. I Crania
anomala. Bib. Zool. 40, pp. 66, 7 pi.
Gratiolet, P.
1857. Etudes Anatomique sur la Terebratule .Australe. Jour, de Con-
cologie. vol. 6, 2 e ser. 2, pp. 209-258. 1 pi.
82 Journal of Entomology and Zoology
18G0. Etudes Anatomiques sur la Lingule anatino. Jour, de Concoloffie,
vol. 8, 2 me. ser. H, pp. 49-107, 129-172, pi. 6-9.
Hancock, A.
18.57. On the Organization of the Brachiopoda. .Anni. .Majr. Nat. Hi.'it.
2 e ser. vol. 22, pp. 141-147.
1859. On the Organization of the Branchiopoda. Phil. Trans. Rov. Soc.
London, vol. 148, pp. 791-S69, pi. .52-66.
Heath, A.
1889. Notes on a tract of modified ectoderm in Crania anoniala and
Lingula anatina. Proc. Biol. Soc. Liverpool, vol. 2. pp. 9.5-104,
pi. 3-5.
Huxley, T.
1854. Contributions to the Anatomy of Brachiopoda. Proc. Roy. Soc.
London, vo. 7, pp. 106-117.
Joubin, L.
1886. Recherches sur I'Anatomic des Branchiopodes inarticules. .\rch.
de zool. exper. 2 e ser. t. 4, pp. 161-303, pi. 7-15, figs. 1-3.
1892. Recherches sur I'Anatomie de Waldheimia venosa Sol. Memm. soc.
zool. de France, t. 5, pp. 554-583, figs. 1-26.
Schulgin, M. A.
1885. Argiope Kovalevskii, Kin Beitrag zur kenntniss dcr Brachiopoden.
Zeit. f. wiss. zool. Bd. 41, pp. 116-141, 2 pi.
Shipley, E. A.
1883. On the Structure of .Argiope. Mittheil. d. zool. Sta. Neap. vol. 4,
pp. 494-520, pi. 39-40.
Sollas, W. H.
1887. Coecal processes of shell of Brachiopods. Proc. Rov. Soc. Dubliru
vol. 5, pp. 318-320, fig. 1.
Vogt, C.
Anatomic der Lingula anatina. Nem. Denkechr. d. Schweiz. Gcsoll.
F. Naturn. Bd. 7, pp. 18, 2 pi.
Delage, Y. et Herouard, E.
1897. Traite Zoologie Concrete. T. 5, pp. 271-2, and pp. 311-12, fig. 442,
pi. 37, 38, 44.
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Journal of
Entomology and Zoology
VOLUME XV, 1923
ITBLISIIEI) QrARTKHI.Y IJY
POMONA COLLEGE DEPARTMENT of ZOOLOGY
CLAREMONT, CALIFOKXIA. V. s. A.
CONTENTS OF VOL. XV
V<iluni<> X\', X umber 1
Merrill, Ida; S«lu>onovor, E.
A Model of the Nasal Chambers
of a White Mouse at Birth, 1.
CliamlM'ilin, R. V.
North American Species of
Mimetus, 3.
Hilton, W. A.
The Nervous System and Sense
Organs, XII, 11.
>(>linii<- X\, \iinil)i-i- il
llillon, \V. A.
Nervous System and Sense Or-
gans, XIII. 17.
Cole, F. R.
Notes on the Early Stages of
the Syrphid Genus Microdin
(Dipteral, 13.
Cole, K. R.
Notes on California Bombyliidae
with Descriptions of New-
Species, 21.
Miiriinon, Sarah
Notes on the Color Changes of
Frogs, 27.
VoluiiK- .W, Niinilier ;{
MwkIows, Donald C.
Notes on the Lepidoptera of
Southern California No. 1. 33.
no<Id.s, Clifford T.
A List of Coleoptera Collected
on the Beach During the
Summer of 1921 at Laguna
Beach, 35.
Campbell, Arthur S.
Some Common Chinese Mol-
hisca, 37.
llillon, \V. A.
The Nervous System and Sense
Organs, XIV, 43.
\ Olunie .W, Nunilier 4
Dodds, ( lifford T.
A New Salt Marsh Mealv Bup.
57.
Campbell, Roy E.
Notes on the Life History of
Dinaparte Wrightii Horn. Gl.
llillon, W. A.
Nervous System and Sense Or-
gans XIV Cent., G7.
INDEX TO VOL. XV
Bombyliidae, 21
Campbell. A. S., 37
Campbell, R. E., 61
Chambevlin, R. V., 3
Cole, F. R., 19, 21
Coleoptera, 37, 61
Color Change, 27
Dinaparte. 61
Diptera, 19, 21
Dodds, C. T., 35. 57
Frogs, 2 7
Hilton, W. A.. 11. 17
Lepidoptera, 33
Marimon, S., 27
Meadows, D., 33
Mealy Bug. 57
Merrill, I., 1
Microdon. 19
Mlmetus, 3
Model, 1
MoUusca, 37
Mouse. 1
Nasal Chamber, 1
Nervous System. 11. 17, 43. 67
Schoonover. E, 1
Sense Organs, 11, 17. 43, 67
Spiders, 3
Syrphid, 1!)
I
(^ NOV 17 1939 i^
VOLUME FIFTEEN NUMBER ONE
i JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
MARCH, 1923
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT o/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
A Model of the Nasal Chamber of a White Mouse at Birth —
Ida Merrill, E. Schoonover 1
North American Species of Mimetus — R. /'. Chnmbcrlin 3
The Nervous System and Sense Organs, XII — //". .7. Hilton. ... H
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A Model of the Nasal Chamber of a
White Mouse at Birth
Ida Merrill and Eugenia Schoonover
Two models were made by the blotting paper method. In one
of these the outer portion of the epithelium was taken as the outer
limit and the lining of the cavity as the other. The other model
was built from the plates which were cut from the interior of the
Upper figure model of the cavity, from outside.
Lower figure model of epithelium from outside.
Both figures X40.
Journal of Kiitoniolofi) and Zoolog\
model. The larger model gives a picture of the mucous membrane
and the nasal chambers, the smaller shows the shape of the nasal
chamber. The knob on the inner side of the larger model is Jacob-
son's organ. The drawings are by Elizabeth Keyes.
(Contribution from Zoological Laboratory of Pomona College.)
Upper figure section of the model of epithelium.
Lower figure model of epithelium from the outside. Jacobson's organ, the
little elevation in the lower t-enttr. X40.
The North American Species of Mimetus
By Ralph V. Chamberlin
In his "Araneides of the United States," Hentz describes three
species under the genus Mimetus : namely, interfector, tuberosus
and syllepsicu.-^. Of these three sijllepxirii:^ has not since been defi-
nitely identified, while tuberosuii is generally regarded as a synonym
of interfector, a disposition with which no fault can be found. In
1882 Emerton described a male from Connecticut under the name
M. ejieiroides: but the practice in recent years has been to refer all
individuals of the genus found in the United States to one species,
interfector, and, accordingly, in current catalogues epeiroides has
been placed in synonymy with that species.
However, a careful study of ample material of Mimetus from
various parts of the country reveals that there are at least five
clearly distinct species that have been confused under the name
interfector. One species occurs on the Pacific Coast apparently
from Washington to southern California and eastward to Texas.
Two species occur in the Northeastern States, the commoner of
these ranging southward as far as northern Georgia. The other
two species are common in the Southern States ; and one of them is
found as far northward as Long Island, N. Y. It seems reasonably
certain that it was one of these two southern forms that was de-
scribed by Hentz as interfector. the particular one being fixed, it is
believed, by the figure of the palpus as indicated later in the notes
on the species. Tuberosus is left as a synonym of interfector; but
syllepsicus cannot be placed at present and is apparently different
from any of the five species here listed.
The males of these five species are easily recognized by the
characters presented in the palpus, the armature of the ectal mar-
gin of the cymbium providing a convenient index. Another readily
observed character of diagnostic importance occurs in the ter-
minal portion of the bulb which in the retracted organ lies adjacent
to the base of the embolus and presents typically two flat or lamellar
lobes projecting proximad. In one species (notius) one of these
lamellar lobes is aborted and in another (puritamis) the second lobe
is itself partly divided or bilobed.
The four species of which females are known may be sepa-
rated in that sex by the characters of the epigynum, which is in the
form of a strongly chitinized, transversely furrowed, caudally pro-
jecting lobe. At the caudal end of the epigynum. or near it on
its dorsal side, is an opening or pit and cephalad or proximad of
this on the dorsal side is a separately chitinized median longitudinal
piece or strip. The position and form of this epigynal opening or
pit, the size and position of caudal end of the median dorsal strip,
4 Journal oi ICntoimildti;) ami Ztioloj:)
and the form and position of the spermathecae as revealed in
cleared specimens furnish good diagnostic characters.
Adults of the species here listed may be identified by means of
ihe following keys in conjunction with the accompanying figures of
palpi and epigyna.
Key to Males
a. Ectal margin of cymbium of palpus with no chitinous, si)init'orni
process proximad of the curved or bent apical one.
b. Ectal border of cymbium with an elevated and sharply limited
lobe at caudal end of scabrous portion of margin, the .surface
of the lobe covered with minute prickles; l)ulb with two apical
lamellar lobe.s (Fig. 5) M. i)tt('rfect(>r Hentz
b.' Scabrous portion of ectal margin of cymbium not ending
caudally in any such sharply defined lobe; apical portion of
bull) bearing only one developed lamellar lobe, the ectal one
being aborted and at most represented by a minute tooth
(Fig. 4) M. iKitiuf! sp. nov.
a.' Ectal margin of cymbium with one or two chitinous processes
or spines proximad of the apical one.
b. With only one spine on margin of cymbium proximad of the
apical one, this toward the base; border scabrous from
apical to basal .spine (Fig. 3) M. fpciroides Emerton
b.' With two spines on ectal border of cymbium proximad of
the apical one of which the more distal one is sometimes
weak; margin scabrous only from apical spine to the more
distal marginal one.
c. Proximal marginal spine contiguous, or nearly so, with
basal lobe or auricle of cymbium ; apical portion of bulb
with neith(>r lamellar lobe at all subdivided or pre.sent-
ing processes (Fig. 2) M. Jicsjtcnis sp. nov.
c' Proximal marginal spine well removed from basal lobe
of cymbium; ai)ical portion of bulb with the larger,
more mesal, lamellar lobe partly subdivided, being
extended at its mesodistal corner (Fig. 1)
.1/. piiritauiiK sp. nov.
Key to Females.
(I. The opening or pit located at extreme caudal end of ei>igynum
and visible in ventral view, the end in this view appearing
notched at the middle: median dorsal strii) extending nearly to
caudal end of epigynum M. i)iirit(nii(s sp. nov.
a.' The pit is on the dorsal surface just proximad of caudal end of
Pomon.i College, Claremoiit, California 5
epigyniim and thus not visible from below, the end not appear-
ing notched at middle ; dorsal strip ending considerably proxi-
mad of end of epigyniim.
h. Opening with no tooth or process from each lateral margin,
not thus partially subdivided; spermathecae essentially
longitudinal ; dorsal strip broader (Fig. 10)
M. notius sp. nov.
/).' Opening partly divided into a distal and proximal portion
by lateral processes; caudal portion of spermathecae bent
at right angles, a distinct enlarged anterior and porterior
portion being connected by a narrower isthmus; dorsal
strip narrower.
c. Lateral margins of epigynum not indented ; isthmus of
of spermathecae narrower, curved, concave ectally.
M. interfector Hentz
c.' Lateral margins of epigynum indented near level of
caudal ends of spermathecae ; isthmus of spermathecae
thick, straight (Figs 7 and 8) . . . .M. hcspei-us sp. nov.
Mimetiis hesperus sp. nov.
In the male of this species the ectal margin of the cymbium of
the palpus bears two conspicuous black spines proximad of the
apical curved one as in piiritanus; but in the present species the
more proximal of these spines is in the re-entrant angle above
basal lobe or auricle, whereas it is distinctly distad of this position
in puritanus. A i-eadily noted difference in the bulb is that the
larger lobe at apex of bulb is entire in hesperus, with no separate
process from inner distal corner as in the eastern form ; and be-
tween this lobe and the conductor there are two folds of conical out-
line not present in the latter species (Cf. fig 2). The female
differs conspicuously in not having the epigynal opening ter-
minal and thus producing a median notch when viewed from below.
The epigynum in its structure most resembles that of interfector,
but differs in outline and in the form of the spermathecae (Cf.
figs. 7 and 8).
Tvpe Localitv. — California : Claremont. Tvpe, a male, M. C. Z.
No. 530.
Other Localities. — California : Stanford : Washington : Camp
Umatilla; Utah; Texas: San Antonio, Austin.
Mimetus puritanus sp. nov.
Mimetus interfector Emerton (nee. Hentz), Trans. Conn. Acad.
Sci., 1882, 6, p. 16, pi. 3, fig. 3.
6 jciuriial lit Kiitomologv and Zoolof^v
Mimetus interfector Keyserling (in part, including those fig-
ured), Spinnen Amerikas, Theridiidae 2, 1886, p. 6, pi, 11, fig. 137.
Thi.s .^^pecies i.s in the female .^ex at once di.';tinguishal)le from
all the others in having the ei)igynal pit at the caudal apex and
visible as a median notch from below (Fig. 6). The male may be
separated from the other species occurring in the eastern and
.southern States by the presence of two subapical spines on the ectal
margin of the cymbium; and from the western hesperus, as indi-
cated above, by the position of the more proximal of these spines
and the form of the larger lamellar lobe of the bulb, which is unlike
that of any other species (Fig. 1).
Type Locality. — New York: Ithaca. Type, M. C. Z. No. 585,
a male.
Other Localities. — New York: Long Lsland, Sea Cliff: Maine:
Ogun(iuit; Mass.: Ipswich, Plymouth; Conn.: New Haven; Vir-
ginia: (Jreat Falls, Falls Church; Georgia: Thompson's Mills.
Mimetus cpeh'oidcs Emerton
Trans. Conn. Acad. Sci., 1882, 6, p. 17, pi. 3, fig. 4.
Known only from the male which is clearly distinct from the
other species in characters of the palpus. In this the ectal margin
of the cymbium possesses a single sjiine toward the basal lot)e, in
distinction from the two preceding species in which there are two
spines on the margin, and from the two following ones in which
there is no marginal spine jn-oximad of the distal one. The ectal
border is scabrous over its entire length from apex to basal spine.
The terminal portion of bull) bears two lamellar lobes, both of which
are simple.
Type Locality. — Mass.: E.ssex.
Immature specimens referred to this species have also been
taken by Mr. Emerton at other places in eastern Massachusett-^ and
at Providence, Rhode Island.
Mi met us interfector Hentz
Journ. Boston Soc. Nat. Hist., 1850, y, p. 3, pi. 4, fig. 12, 13.
Mimetus tuberosus Hentz, ibid., p. 3, pi. 4, fig. 14.
Of each of the two species of Mimetus occurring commonly in
the southern States, individuals may be found which match the fig-
ures of interfector given by Hentz reasonal)ly well. I believe the
species to be fixed, however, by the figure of the palpus of the male
which, in spite of its general inadefpiacy, shows two prominent
lobes ))i-o.jecting jiroximad fi'om the bulb that are ai)i)art'ntly the
two lamellar lobes present in the one species, whereas in the other
species, listed below as M. notius, sp. nov., there is but a single
I
I
PomoiKi Ccilley;e, Clarcnioiit. California 7
lamellar lobe. In the species thus considered to be fixed as the
true interfecior of Hentz the ectal margin of the cymbium lacks
spines; the scabrous border ends proximally abruptly in a lobe
elevated above the general surface and on which the area of
prickles is broader, a very characteristic feature enabling one to
detect the species at a glance (Fig. 5). The form of the opening
of the epigynal pit is similar to that of ]iesperus, being partly sub-
divided by a projection from each lateral margin and thus differing
from that of notiic^. The spermathecae also present a caudal and
an anterior larger lobe connected by a narrower, weakly curved,
isthmus.
Type Locality. — Alabama.
Other Localities. — Alabama : Morgan, Birmingham ; Georgia :
Atlanta; Louisiana: Shreveport, Covington, Shrewsbury; North
Carolina; New York: Sea Cliff.
Mimetua notii(.!<, sp. nov.
In this species the opening of the epigynum lacks projections
from its lateral margins, and the median dorsal strip is broader and
more conspicuous than, e. g., in M. interfector or M. hesperus; the
spermathecae are essentially longitudinal as shown in fig. 10. The
male differs from all the others here considered in having on the
distal portion of bulb of palpus only a single lamellar lobe, the ectal
one being absent or represented only by a slight tooth at base of the
developed lobe. The ectal margin of the cymbium lacks spines
proximad of the apex and its scabrous border runs out gradually,
not ending in any such abruptly elevated lobe as occurs in intvi-
fector.
Type Locality— Runnymede. Type, a male, M. C. Z. No. 551.
Other Localities. — Florida: Altoona, Daytona; Louisiana:
Shreveport. Mansura; North Carolina: Raleigh.
Fig. 1. MimetuN piiritatiiis sp. nov. Rijirlit paljiiis of malo,
subectal view. 2. M. Iirspmis .sp. iiov. Ri^ht iiaiinis of malo,
.similar view. .'5. M. tpciraidcs Emerton. Ri^ht i)aipiis of malo
(type) from a more (ior.sai aspect, the hematodocha (iisteiuied. 4.
M. notius sp. nov. Right i)alpu,s of male, subectal aspect. 5.
M. Interfector Heiitz. Right palpus of male, subectal aspect.
Fig. 6. Mimetiis puritanus sp. nov. Epigynum, ventral
view. 7. M. hesperus sp. nov. Epigynum, ventral view. 8. M.
hespei-u^ sp. nov. Epigynum viewed from above by transmitted
light, showing opening, dorsal strip, and the spermathecae in sil-
houette. 9. M. interffctor Hentz. Epigynum in ventral view.
10. M. notius sp. nov. Epigynum viewed from above by trans-
mitted light to show form of opening and of dorsal strip and the
spermathecae in silhouette.
XII. Enteropneusta
For our general knowledge of the central nervous system of
this group we have the papers of Spengel, 1884-1894, Bateson, 1886.
Of the development of the nervous system and the larvae the
work began in 1870 with the study of the so-called Tornaria larvae.
Bateson, 1884-5, worked out the life history of a Balanoglossus
form and later Spengel, '94 and Morgan, '91 and '94 gave an ex-
tended account of the Tornarian forms, including a good account of
the nervous system. Ritter, '94 and Davis, '08, gave stages in the
development of Tornaria and DoUchoglussus, and Herder, 1909, also
gave an account of development in which the nervous system was
included.
In various accounts of the position and structure of the nervous
system especially as summarized in text books and other places,
there seems at times to be some difference in the descriptions but
I think for the most part the differences are in the way of express-
ing much the same idea so that no real difference is introduced.
In all cases the nervous system is as a whole epidermal much
as in Phoronis and in starfish. The epithelium everywhere is more
or less made up of columnar cells at the surface with a deeper
nervous layer of fibers, in part branches from the surface cells, and
a few deeper cells. In places the epidermic nervous system is
more marked. The whole body then might be described as covered
■with a plexus of nerve cells and fibers ; the thicker parts of the
plexus in places form the so-called nerves. The chief nerves of this
sort are a dorsal and ventral tract in the body region below the
collar with a circular band connecting these at the lower edge of
the collar, and a concentration of fibres about the base of the pro-
boscis, but the greater concentrations are in the collar itself. In
the dorsal and ventral surfaces of the collar just under the epi-
dermis is a concentration of nerve cells and fibers but the chief and
central concentration of nervous tissues is in the form of a thicker
cord running through the cavity of the collar on the dorsal side,
although connected with the epidermis at each end. This central
nervous system is continuous with the proboscis thickening in front
and as just described, with the dorsal and circular nerve tracts
behind.
To summarize, the nervous system may be described as fol-
lows:
1. General epidermal plexus continuous with other parts.
2. Basal proboscis ring continued into the proboscis by a
more diffuse band.
3. Ventral body nerve continued into ventral collar as a thin
layer.
Fijr. 2."). DiaKranis to show the position of the nervous system in Dolicho-
glossus. Nervous system shown by heavy lines below the surface.
1. Longitudinal section. 2. Cross section throuRh the proboscis.
3. Central portion much enlarged. 4. Another part of the surface.
.'). Neural epithilium much enlarged.
PoiTioiiii College. Cl:irciii(int. California
13
4. Dorsal collar nerve somewhat cut off from the two follow-
ing.
5. Dorsal proboscis nerve continued above.
6. Central nervous system running through the central region
of the collar on the dorsal side and continuous above with the pro-
boscis nerves and below the collar with the dorsal body nerve.
The dorsal nerve of the collar and the thick central nervous
system of the collar are more or less joined by the strands and
they together make a sort of nerve tube thin on the dorsal side but
thick below.
The histological structure of the nervous system reveals be-
sides the usual epithelial cells of the surface, bipolar supportive
FJK. 24. Nervous System and Sense Oi-Rans of Enteropneusta. A. I)iaKi-ani
of Bo?OHO(//o.ss)(s showing position of the nervous system. B. Anoth-
er diagram of Balaiinglossus in sectional view. Spengel. C, D.
Sections of developing nervous system. Morgan. E. Larva after
Herder. F. Apical eye of tornaria larva. Spengel. G. Eyes of
Tornaria after Morgan. H. Eye of Tornaria after Spengel.
14 Ji)urn,il of F,ntiiniiil()f:\ and Zonlo^jy
cells reaching from the surface to the depths of the nervous system
and also probably bipolar sense cells as well as more or less deeply
placed multipolar nerve cells giving off fibers to the nerve areas.
The epidermis is a general organ of sense, the exact nature of
which has not been very clearly determined. Spengel considers
that about the proboscis in its ventral face there are points espe-
cially sensitive. In fact at this place he describes a deep depression
which he regarded as a special sense organ.
In the larval stage the first suggestion of a nervous system we
find in the development of the apical plate which in later stages
develops eye spots as simple caps of ectodermal cells surrounded by
pigment. The eye spots become anterior in position with a pocket
of the clear cells each ending in a point. Between the two eye-
cups a mass of pigment develops. At the base of the apical plate
nerve fibers begin to be seen.
At metamorphosis in a region where the collar will develop a
transverse groove forms near the mid-dorsal line. In the mid-
dorsal region a strip of ectoderm not crossed by grooves makes the
beginning of the neural plate. It sinks beneath the surface and
folds of the adjacent ectoderm or neural folds meet over it, and in
this way the neural tube is formed.
BIBL10GR.\PHY
Bateson, W.
18?4. The Early Stages of the Development of Balanojrlossus. Q. Jour,
mic. sc, vol. 24, pp. 207-2.3.1, pi. 18-21.
188."). Later Stages in the Development of Balanoglossus kovalevskyi.
With a suggestion of the .Affinities of Enteropneusta. Q. Jour,
mic. sc. vol. 2.5, supp. pp. 81-122, pi. 4-9.
1886. Continued account of the Later Stages in the Development of
Balanoglossus kovalevskyi, and the Morphology of the Enterop-
neusta. Q. Jour. mic. sc. vol. 26, pp. 512-533, pi. 28-33.
1886. On the Morphology of the Enteropneusta. Stud. M. Z. vol. 3. pp.
37-65, pi. 7-12.
Davis, B. M.
1908. The early Life History of Dolichoglossus pusillus. Univ. Calif.
Pub. zool. vol. 4, no. 3.
DawydoflF, C.
1909. Beobachtungen uber den Regenerationsprozess bei <len Rnterop-
neu.sten. Zeit. f. wiss. zool. Bd. 93, pp. 237-305, pi. 13-16, 23 text
fig. P. 293 N. Syst.
Haldeman. G. B.
1886. Notes on Tornaria and Balanoglossus. J. Hopkins Univ. Circ. vol.
6, pp. 44-5.
Herder, K.
1909. Zur Entwickelung von Balanoglossus clnvigerus. Zool. aiiz. vol.
34.
PoiiKiiia Collff^c-. Claremont, California 15
Hilton, W. A.
1919. The Central Nervous System of Dolichoplossus. Jour. Ent. Zool.
vol. 9, pp. 59-61, figs. 1-5.'
Kovalevsky, A.
1866. Anatomie des Balanoglossus (Delle Chiaje). Mem. .\ca(l. inp. se.
St. Petersburg. 7e ser. vol. 10, no. 3, :i pi.
Koehler, R.
1886. Contribution a I'etude des Enteropneustes. Recherch anat. sur Ic
Balanoglossus sarniensis nov. sp. Internal. Monats. Anat. Hist.
vol. .3, pp. 1.39-190, pi. 4-6.
Metchnikov. E.
1870. Untersuchungen ueber die Metamorphose einiger Seenthiere. 1.
Ueber Tornaria. Zeit. f. wiss. zool. vol. 20, pp. 131-144, pi. 13.
Morgan, T. H.
1891. The Growth and Development of Tornaria. Jour. Morph. vol. 5
pp. 407-458, pi. 24-28.
1894. The Development of Balanoglossus. Jour. Morph. vol. 9, pp. 1-86,
pi. 1-6.
Ritter, W. E.
1894. On a New Balanoglossus and its Possession of an Endostyle
Zool. anz. vol. 17, pp. 24-30.
1900. The Movements of Enteropneusta and the mechanism by which
these are accomplished. Biol. Lect. Woods Hole. vol. 3.
Spengel, J. W.
1884. Zur Anatomie des Balanoglossus. Mitt. zool. St. Neap. vol. 5, pp.
494-508, pi. 30.
1893. Die Enteropneusten des Golfes von Neapel und der angrenzenden
Meeres-Abschnitte. Fauna Flora Golfes Neap. 18 Monog. 758 pp.
37 pi.
Spengel, J. W.
1877. Ueber den Bau und die Entwickelung des Balanoglossus. Amtl.
Ber. 50. vers. d. Naturf. u. artze. Munchen.
1903. Neue Beitrage zur Kenntniss der Enteropneusten. 1. Ptychodera
fluva Esch. zool. Jahrb. Bd. 18, pp. 271-326, pi. 24-29, 5 text figs.
1904. 3. Fine neue Enteropneusten art aus den Golf, von Neapel. Zool.
Jahrb. vol. 21, pp. 15-362, pi. 20-22, 10 text figs.
Stiasny, G.
1914. Studien ueber die Entwickelung des Balanoglossus clavigcrus
Delle Cheaje. 11 Darstellung der weiteren Entwickelung bis zur
metamorphose. Mitt. zool. St. Neap. vol. 22, pp. 255-290, pi. 59.
13 text figs.
1914. Studien ueber die Entwickelung des Balanoglossus clavigerus Delle
Chea'e. I Die Eentwickelung der Tornaria. Zeit. f. wiss. zool.
vol. 110, pp. .36-75, pi. 4-6, 24 text figs.
Ii^ NOV 17 1939 ^\
VOLUME FIFTEEN NUMBER TWO
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
JUNE, 1923
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT o/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
Nervous System axd Sense Orcaxs, XIII — ff\ A. Hil/on 17
Notes on the Early Stages of the Syrphid Genus Microdin
(Diptera)— T. R. Cole 19
Notes on California Bombyludae with Descriptions of New-
Species— T. R. Cole 21
Notes on the Color Changes of Frogs — Sarah Mnrimon 27
Entered Claremoiit, Cal.. Post-Office Oct. 1. isio. as second-clasa matter, uijder Ait ul Coiigrtsb of
Vlarcb 3. ISTU
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XIII. Cephalodiscus and Rhabdopleura
Cephalodiscus. The first report on this animal including a
sketch of its anatomy was by Mcintosh, 1887, and later by Lang,
1890, and Harmer. Delage and Herouard, 1897, summarize the
knowledge of the nervous system about as follows :
The nervous system is a thickening of the epidermis on the
dorsal surfaces of the tentacles. The histological nature of the
nervous system was a little studied, but cells and fibers under the
epithelium as in echinoderms were described.
Mastermann, 1903, describes the central ganglion over the sub-
neural blood sinus ; its position is as in Actinotrocha. This gives
off below a pair of large nerves each of which divides into six
branches for the six pairs of tentacle arms. Above it is prolonged
^#^""
Fig. 26. A. Cephalodiscus showing location of the nervous system after
Hammer. B. Cephalodiscus after Masterman. C. Rhapdopleitm
showing position of nervous system after Delage and Herouard.
18 Journal oi Kiitomology and Zoolnpi'
into two large branches which follow the dorsal line of the epistome.
Laterally from the ganglion two other nerve l)ranches go to the
epistomal disc. On the ventral surface of the trunk is a medial
longitudinal band which is continued into the peduncle. According
to Mastermann the mid-dorsal and two lateral epistomal branches
have homlogues in Balanglossus and Actinotrocha.
Rhabdopleura. The account of the structure of Rhabdopleura
which is usually given is that of Fowler, 1892. Other accounts
which however give little of the nervous system are those of Allman,
1869. and I.ankester, 1874.
The central nervous system is represented by a thickening of
the ectoderm in the median region of the neck below the nucal pore
between the branches of the tentacles. There is a differentiation
of nervous tissue much as in Rhahdoplvura or Bahninglossiu^. A
black pigment spot is located at the tiii of the preoral lobe and may
be an eye-spot,
BIHLIOGRAI'IiY
Fowler, G. H.
1892. The Morphology of Rhabdopleura normani Allm. Festachrift.
zur 70 Geburtst. R. Leuckart'.s Upzig. pp. 293-297. pi. 30.
Harmer, S. F.
1887. Cephalodiscu.s. Zool. of H. M. .S. Challenger, vo. 22, part <)■'!. pp.
.39-47, 2 fip.s, 7 pi.
Lanp, A.
1890. Zum Vor.staiulniss dor Organisation von Cuphalodiscus dodecalo-
phu.s Mcintosh. Jen. Zeit. f. Nat. vol. 25, pp. 1-12.
Lankester, E. R.
1874. Remarks on the Affinities of Rhabdopleura. Q. .lour. niic. sc. vol.
14, pp. 77-81.
18F4. Contribution to the KnowledKc of Rhabdopleura. Q. Jour. niic. sc.
vol. 24, pp. 622-647, .5 pi.
Mastermann, A. T.
1897. On the Diplochorda. 1. Actinotrocha. 2. Cephalodiscus. Q. Jour,
mic. sc. vol. 40, pp. 281-.3C6, pi. 18-26.
1903. On the Diplochorda. 4. On the central complex of Cephalodiscus.
Med. Q. Jour. mic. sc. Ns. vol. 46, pp. 32-33.
Schepotief, A.
1908. Cephalodiscus dodecaloptuis. Zool. Jahrb. anat. vol. 2h, pp. 40.")-
494.
Notes on the Early Stages of the Syrphid
Genus Microdon (Diptera)
By Frank R. Cole, Stanford University
The peculiar larvae of the Syrphid flies of the genus Microdon
have been described by several entomologists, but they are known
in only a few species. Enthusiasts in past years placed these
bizarre forms among the molluscs in two or three instances, and
one entomologist stated that they were the early stages of a Coccid
on oaks. , Wheeler has given a vei'y interesting account of some
of these early stages and the habits of the flies. In America the
larvae are recorded only from ants nests, but Wasmann states that
they may be found in the nests of certain wasps and termites.
The.v live in nests in the soil, under rocks or under the bark of
old logs.
The larvae creep very slowly, with a wave-like m_otion of the
flat ventral sole, which is fringed and applied closely to the sur-
face over which they are travelling. Their food is probably, as
Wheeler believes, the minute pellets of food ejected from the
hypopharyngeal pockets of worker ants after the moisture has been
extracted. There is evidently one brood in a year, the flies emerg-
ing in May and June.
In May, 1917, the writer found a number of pupae of Micro-
don cothurnatus Bigot, while collecting in the Hood River Valley
of Oregon. The type of this species came from "Mt. Hood," prob-
ably somewhere in the valley north of the mountain. While pull-
ing off the bark from an old pine log an ant's nest was uncovered,
and among the frenzied inhabitants of the nest a number of
Microdon pupae were noticed. The ant was later determined as a
subspecies of Camponotus maculatus. At this date. May 19, there
were no larvae of the fly present and the pupae were all full.v
developed. Eighteen pupae were taken, most of them rather closely
crowded near the entrance to the nest ; all around them were empty
puparia, bearing evidence that the nest had been used for several
seasons by the flies. There were several adult flies around the log,
some of them freshly emerged, but the ants were so aroused at the
disturbing of their domestic tranquility that they quickly drove out
any strange insect that came near. The puparia taken were allowed
to become too dry and only two adults emerged out of the lot.
In April, 1921, some observations were made on another
species of Microdon. A student at Stanford University, Mr. Her-
bert Mason, found a single larva in a nest of Camponotus maculatus
vicinus Mayr. This specimen was reared by Mr. Carl Duncan and
the specimen and notes regarding it kindly turned over to the
writer. The species proved to be Microdon pipcri Knab, a beautiful
dark blue species which ranges north along the Pacific coast region.
The larva was not closely examined by the writer, but in the
notes made on the specimen' the color was given as largely pale
20
Journal of Entomologx an<l Zoolojry
bluish green, with median ridge and the margins of the body
brown. The median ridge was quite prominent in the larva. The
coarse reticulum on the body has a pattern somewhat like that
figured for M. tristis, as can be seen from the figure. The length
was 11 mm.
The puparium shorten.s to about 9 mm. The reticulum is much
more distinct than in the larva and two prothoracic tubercles push
out (in the specimen described one of the tubercles did not push
through the body wall). The reticulation is arranged in a more or
less symmetrical design and when examined under a high magni-
fication is seen to be made u]5 of two types of processes; those on
the dorsal ridge and along the sides just above the fringe are of a
shape which might be likened to an inverted wine glass and the
other processes are (luite short and composed almost entirely of the
white stalked portion (see figs. 6 and 7). The base in both cases is
dark brown and the apical portion white. From above the boJv
appears to be covered with white di.scs arranged in a reticulated
pattern, the center of each disc with a depression and a minuU'
cavity which appears to penetrate almost or quite through the
body wall. These minute structures may function as pores. The
anterior margin of the ventral fringe of the body is deeply notched
in the middle as shown in figure 4. The structure of the marginal
fringe is shown enlarged in figure 8. The fly emerges from the
puparium by bi-eaking off the cover in three rather symmetrical
pieces, illustrated in figure 2.
The specimen described was taken the last of March and soon
commenced to pupate, the puparium being fully colored by April 8.
The adult emerged just a month later.
Wheeler notes that the most typical and frequent hosts of t'
Microdon larvae are ants of the genus Fonn'uct but Wasmann has
recorded a speci<"< "•'""•" '-<../'.///, iH \!.-.il,iir;,<, ■,,• -.^ a host.
%^
2 •* 8
Fijr. 1- F'uparium of Microdon piprii Knab; 2, anterior portion of puparium,
showing symmetrical breaking; 3. posterior respiratory tubercle;
4, mari;inal frinpe of puparium, showing split in anterior region ;
b, reticulations of two kinds, those with a short, and those with a
high base; 6, and 7, portions of the reticulations more highly mag-
nified; 8, marginal fringe, greatly magnified.
Notes on California Bombyliidae with
Descriptions of New Species
Frank R. Cole, Stanford University, Cal.
The sun-loving Bombyliidae have always been a favorite group
with the writer, as the rather abraded specimens in his earliest
collections will bear evidence. California is rich in species of these
flies and notes on a few of the interesting forms are given below.
During the past two summers the writer has spent some time
in Mill Creek Canyon in San Bernardino County. Paracosmus
morrisoni 0. S. is a very common form in this locality and is
usually taken alon<T roads and paths in the bright sunlight. Aphoe-
bautu!^ vittatufi Coq., a trim, beautifully marked little species,
occurs along with the above, but is not so common and is often
harder to catch. Villa squamigera Coq. and Villa mira Coq. are
not uncommon in the Mill Creek region, the latter species more
abundant in August, when it is found out in the sandy river
washes. Villa miscella Coq. is seldom seen and is quite wary,
flying up and down sandy roads for long distances when disturbed.
In Glen Martin, in this same general region but at a higher alti-
tude, one occasionally finds Rbabdoselaphus setosa Cresson, a little
species with a very long proboscis ; it is usually taken on the wing
in the middle of the day, hovering near the ground. With the first
days of autumn specimens of Villa autumnalis Cole begin to ap-
]5ear, frequenting the yellow flowers of Ericameria and Chry-
sotliamnus, and now and then a specimen of the beautiful golden
Lordotus diversus Coq.
Villa chromolcpida new species.
Female. Length 7 mm. Black, clothed with bright iridescent
scales; front tibiae without bristles; wings hyaline.
Head rather large in proportion to the body ; proboscis pointed
and scarcely projecting beyond the oral margin; palpi small, black,
cylindrical and black pilose. Antennae black, first joint about
twice as long as second and with black pile; third joint twice as
long as first two combined and gradually tapering toward apex
(see fig. 9), the apical bristle minute. Frons shining black, with
erect black pile and sparse golden scales which are purple in
cei-tain lights. Face projecting (see fig. 10), shining black, with
scales like frons but denser, pile "short, black, reclinate. Cheeks
shining black, bare of pile or scales. Occiput black, densely
clothed with scales like those on face and frons ; next the hollowed
out back of the head a line of short, fine, yellowish pile.
Mesonotum and scutellum shining black, with golden green
scales, purple by reflection; the median portion of dorsum with
22 Journal of F'titomoloK)' and Zoology
erect blackish pile, the front and margins with white pile, stiff a^^ '
erect just back of the head. Pile of scutellum sparse and white.
Pleura .'^hining black, with rather long, dense white pile on the
upper mesoi)leura. the lower part of mesopleura and other pleura'
plates with sparse black pile, not obscuring the ground color; sti V,
blackish bristle-like pile above front coxae. The coxae and pleura
with a few scattered iridescent scales. Stem of halteres yellow, thi-
knob white, with a black mark on anterior margin; tuft of pile
before halteres largely yellow.
Abdomen black, with erect white pile on sides of first and on
anterior corners of second segment ; beyond this the pile is ver
sparse, black,, reclinate and .'Scarcely noticeable. On each side of
lK)sterior margin of fir.^t visilile tergite some scales like those o.i
thorax; on the other abdominal tergites and sternites there is a
dense covering of tomentum or scales, largely colored like those ol
thorax and in a definite design on dorsum ; in the center of eacli
tergite beyond the fir.st vi?ibb one a round spot with sparse bla
scales, on each side a larger oval spot covered with black .scales
which have a inirplish color in some lights ; these lateral spots mis.--
ingon seventh segment, which is almost wholly covered with irides-
cent scales. The venter black, with a wide median portion cloth •
with black tomentum, the sides with iridescent scales as on th -
more or less telescoped, the last two segments scarcely visible;
color of pollen and pile as in male. Apices of femora an ochre
dorsum. Femora and bases of tibiae brownish yellow, the rest of
legs black; all the spines and pile of legs black, front tibiae without
bristles, the anterior tarsi with claws almost as large as on the
other tarsi ; femora with a few yellowish, iridescent scales and
some black ones; tibiae and tarsi with black .scales. Wings hya-
line, iridescent; the costa and veins at base yellowish, toward
posterior margin black; fork of radius rather angular at base.
The epaulets with purplish iridescent .scales.
Holotype, a female, collected in Mill Creek Canyon, Cal., July
20, 1920 (F. R. Cole), in the collection of the California Academy
of Sciences.
The type female is the only specimen known and is not closely
related to any species seen by the writer. In Coquillett's table of
species it would run to mcrccdis. It is distinct from any de.scribed
Mexican species.
Anipliico.-iniKs faiidKZcci new species.
Female. Length 6 mm. A slender species, the body largely
shining black, the legs yellow.
Ui)per two-thirds of frons black, including the large ocellar
tubercle, the lower third yellow; pile sparse, fine, white, the narrow
orbits silvery pollinose. Face short, projecting, the central portion
shining black, sides yellow and with silvery pollen; antennal fovae
Pomona College, Claremoiit. California 23
deep and connected; first antenna! joint slightly longer than
second, yellow; second and third joints black, the third joint about
as long as the first two combined, narrower (see fig. 6), with a
short sub-apical style. Vertex and upper occiput rather full (see
fig. 7), black, the lower occiput and cheeks yellow, occiput largely
silvery pollinose, the pile minute and whitish.
Mesonotum and scutellum shining black, the pile on median
portion of mesonotum and on scutellum short, blackish, on margins
of mesonotum white. Humeral callosities yellow, silvery pollinose;
a silvery pollinose, white pilose spot just back of humeri. Pi-escu-
tellar callosities partly yellow. Pleura shining black, the upper
mesopleura, the metapleura and hypopleura silvery pollinose and
white pilose. Halleres white.
Abdomen largely shining black, rather broad posterior mar-
gins of all segments yellowish ; apical half of seventh visible seg-
ment lemon yellow ; yellow on first segment reaches lateral margins,
on the second to sixth segments it does not do so. Pile of abdomen
very fine, sparse, white, longer on sides of first and second. Venter
largely brownish yellow, blackish at base, lemon yellow on genitalia.
Femora, tibiae, first tarsal joint, apex of fifth and base of claws
honey yellow; third and fourth tarsal joints, apex of second and
base of fifth blackish. Coxae and trochanters colored like femora,
a black spot below on base of trochanters. Wings hyaline, all
veins yellow at base, toward apex and posterior margin blackish.
All cells on posterior margin of wing wide open (see fig. 8).
Holotype, a female, collected at Palm Springs, Cal., May 20,
1917 (E. P. Van Duzee), in collection of California Academy of
Sciences. The type a unique.
This species differs from elegans Coquillett in having the first
antennal joint yellow and in the greater extent of black on the
abdomen. Coquillett gives no structural characters to distinguish
his species. The above described species differs from cincturus
Williston, from Mexico, in the smaller size and in the color of the
antennae and legs, cincturus having entirely black legs.
Metacosmiis nitidus new species.
Female. Length 5.5 mm. Head black, a small amount of yel-
low on sides of oral margin. Ocellar tubercle slightly above
middle of frons but the lower ocellus nearly in the center; upper
half of frons with white pile, the lower part with black; frons
shining black, the narrow orbits silvery pollinose. Antennae black,
rather short and thick, the second joint larger than first (see fig.
4). Upper face and lower frons near base of antennae silvery pol-
linose; face short, shining black, distinctly projecting. Occiput
thinly gray pollinose, short, sparse white pilose; on the under side,
back from mouth opening, an oval yellow spot on each side of
middle. Proboscis not projecting beyond oral margin.
J4 Jnmna! (it F.iitomolojiv and Zoology
Thorax shining black, the dorsum with short, sparse white
pile; scutellum shining black, with short white pile. Humeral
callosities and lower half of pleura gray pollinose. Stem and under
part of knob of halteres blackish, the most of knob white.
Abdomen shining black, finely punctate and with short, sparse
whitish pile; hind margins of visible segments one to four narrowly
yellowish white, i)roader on the first. Abdominal i)ile appears
white in certain lights but is largely dark colored. Sternite of
seventh segment projects downward in a noticeai)le triangle as seen
in profile. Pile around genitalia rather dense and whitish. Ven-
ter black, the hind margins of first five segments yellowish white.
Legs wholly black, the pile fine and short. Wings hyaline, rather
broad and rounded, the veins black and strong; R24-3 curved
slightly forward at tip (see fig. 3).
Holotype, a female, collected at Huntington Lake, Fresno
County, California, 7000 feet, July 15, 1919 (E. P. Van Duzee), in
the collection of the California Academy of Sciences.
Paratype.s. — Two females, taken in the tvi^e localitv, Julv 8.
1919, by Mr. E. P. Van Duzee.
This species is evidently near M. e.r(7(.s Coquillett, but differs
in the color of the legs and in the wing venation. The only other
species in the genus is mancipennis Coquillett an eastern form,
which has the face and the stems of the halteres white.
Acreotrichwi macuUpennitt new species.
A velvety brown species with thickly spotted wings; the pro-
boscis slightly longer than the head.
Male. Length 4.25 mm. Head black, brownish iiollinose, the
face and vertex with rather long and erejt black pile. Occiput
rather flat ; occiput and cheeks with black pile. Oral opening
large, the antennae i)iaced on the uiii)er edge (see fig. 2) ; first and
second anetnnal joints rather slender, the first slightly longer than
the second, the third slightly longer than the fir.st two combined
and considerably widened near the middle, the style short and
subapical (see fig. 2) ; pile on upper side of all antennal joints
black. Proboscis black, projecting twice the length of the antennae
l)eyond the oral margin. Palpi black, very slender, with black
l)iie, projecting beyond oral margin about one-third as far as
proboscis.
Thorax black; mesonotum velvety black, shading to a sepia
brown on the margins; the pile of dorsum erect and yellowish.
ai)pearing brown in certain lights. Scutellum velvety black, with
comi)aratively long, coarse yellowish pile. There are indications of
two median black vittae on the anterior part of the mesonotum,
.separated by a fine brown line. Pleura brown pollino.se, the sparse
Collect-, ClareniDiit, Californi.-i
25
pile on mesa- and sterno-pleura brown. Stem of halteres yellowish,
the knob yellow above and blackish bi-own below.
Abdomen black, sepia brown pollinose, with rather long, erect
yellowish pile, nowhere dense enough to obscure the ground color.
Venter like the dorsum, the pile shorter and more reclinate. Sev-
enth visible segment projecting over the small eighth, the genitalia
quite small, colored like the abdomen, the upper and lower forceps
about equal in size and closing over the internal organs. Knees
reddish, the rest of legs black ; coxae and femora with long black
pile. Wings whitish hyaline, densely maculated with dark gray
and with remarkable thickenings of the membrane, some of which
appear to form supernumerary cross-veins (see fig. 1). The veins
near the posterior margin of the wing are wavy.
EXPLANATION OF PLATE
Fig-. I. Wing of Acreotrichus nutculipetiiiis n. sp. ; fig. 2, head of A. mucn-
liljenuis; fig. 3, wrng oi MetacosiniiK nitidus n. sp. ; fig. 4, head of
M. nitidus; fig. .5, wing of PamcosDii^s vionisoni O. S. ; fig. (5, antennae
and front of head of Amphicosmiis vunditzeei n. sp.; fig. 7, head of
.4. vaudiizeei; fig. 8, wing of A. vunduzeei; 9, antenna of Villa
chrcmohpida n. sp. ; fig. 10, heal of V. cliroviolepida ; fig. 11, head of
Rliabdoselaphus setosiis Cresson.
26 Journal of Entomology and Zoology
Female. In general very much like the male but lighter in
coloration. Pile of cheeks and lower occiput yellowi.sh, on the re.st
of the head and on the antennae reddish brown. Eyes widely
separated, the pollen of frons more buff colored than in male, the
pile shorter. Pollen of mesonotum much lighter in color than in
male, the pile shorter and paler. (Iround color of coxae and pleura
yellowish brown in some specimens, the pile yellow. Knob of
halteres .scarcely darkened below. Abdomen in dried specimens
"yellow, also the tibiae except apices and bases of the four front
tarsi. Pile and fine setulae of femora and tibiae yellowish.
Holotype, a male, and allotype, a female, collected on the sand
dunes near Golden Gate Park, San Francisco, Cal.. September 10.
1920 (F. R. Cole), in the collection of the California Academy of
Sciences.
Paratijpes. — Two specimens in the Cal. Acad, of Sci., taken in
the type locality, and five specimens in the writer's collection, taken
with the types.
In 1895 Cociuillett described Acreutriclixs anuricauiis from a
single male specimen taken in the state of Washington. This litt'c
species has hyaline wings, the antennae are quite different and the
proboscis comparatively longer. In May, 1917, the writer too';
a single male specimen of A. americanus near Hood River, Ore-
gon; it appears to be a rare species. A. atrati(,s Coquillett, from
Mexico, has a slender third antennal joint, three times as long as
the first two combined and of nearly an equal width; the wing^
are grayish hyaline. The three other known species in the genus
are described from Australia.
Notes on the Color Changes of Frocrs
Sarah Marimon
In all these experiments I chose two identical frogs, and kept
one in normal conditions while the other was being subjected to
change.
Tree frogs, Hyla regilla.
I. June 16. Hot water (about 30 to 35 C). Left the frogs
for one hour.
The spots of the frog faded out, giving a lighter appearance.
However the background seemed much the same as the control.
Control. Tap water (about 15-17" C). Spots distinct.
Grayish green frog.
n. June 17. Hot water. The frogs for this experiment had
peculiar red and green markings.
The whole tone was lighter at the end of an hour and one-half.
Spots somewhat more indistinct than at first.
Control. Color unchanged.
III. June 16. Cold water. (Cooled with ice— 2 C.) The
frog was somewhat darker in color : the spots stood out more dis-
tinctly than previously.
Control. Tap water (15-17 C.) Color unchanged, spots
showing distinctly — not so distinctly as those of the frog in cold
water.
IV. June 17. Cold water, a. The frogs were rather light in
color. Darker spots more distinct.
b. Two frogs grayish green in color. The color became
darker, spots more distinct.
V. June 17. 5:00 P.M. Two frogs with red .streaks down
the backs.
One jar covered with green tissue paper, the other left as a
control.
June 18. 10 A. M. Lighter in tone than the control. The
red streak changed to light sandy color. Spots lighter,— greenish
along the sides.
Control. Same as the day before, apparently. Spots dark
grey, grey sides, broad reddish streak down the back.
11 A. M. The frogs reversed.
June 19. Red streak narrower, sandy colored. The whole
cast of the frog was lighter and more greenish.
28 Journal of F,iUoniolo)i\ and Zoology
Streak dark reddish. Frog much darker than the one in green
jar.
VI. June 21. Green and cold. To see which has the greater
effect, the background or the temperature.
a. Two frogs rather light in color.
The spots are more distinct but the whole color is lighter
than the control.
b. Two frogs rather dark in color.
Slightly lighter. The dark colored frogs do not seem to change
as readily as the lighter ones.
These experiments would .seem to indicate the greater effects
of the background. However there was some chance for error
here, because (1) the experiment was only over a period of two
hours, and (2) because the frogs objected to the cold water, and
when they were not watched, they would climb up out of the water
and cling to the side of the jar.
VII. June 17. I put two frogs in a jar lined with leaves.
One frog very reddish, the other grayish green.
June 21. The grayish green frog much greener, lighter in
tone.
The reddish frog much lighter in tone l)ut still decidedly red-
dish in color.
June 23. The red frog still reddish.
The green background lightened it but did not change its color.
VIII. June 17. 5 P.M. Red cover to the bottle. Placed the
frogs in the jars.
June 18. 10 A. M. Slightly darker. Control. Color un-
changed.
June 20. a. About the same shade as the other frog (i. e. the
control) only with a more reddish tinge, b. Distinctly lighter, and
more reddish in color.
June 21. Frogs had each a sandy streak down the back.
The .streak brighter reddish. The whole tone of the frog
slightly darker than the control. The frogs reversed. Streak
sandy colored.
June 22. Streak brightly reddish. Whole tone of frog much
lighter. Streak sandy colored. Darker than the one in red.
VIII. (a) June 17. 5 P. M. Took two greyish frogs with
no particular color showing. Placed one in a jar covered with
yellow tissue paper. The other frog used for a control.
June 18. Much lighter than the control. Seemed to have a
yellowish tinge. Spots faded somewhat.
Reversed the frogs.
Pomnnn CoUciiC. Clarcinont, Caliturnia 2'^
June 19. Lighter, the spots more faded than when in the con-
trol. The difference between the two not so marked as on June 18.
June 20. Slightly lighter, more yellowish in tone. Results
not so distinct.
VIII. b. June 21. Two frogs with reddish streak (June 20) .
The streak more yellowish, now has a distinctly yellowish tone.
Spots lighter. Whole tone more yellow than control.
Reversed the frogs.
June 22. Yellowish in tone. Red streak now very yellowish.
Spots lighter.
IX. June 17. 5 P. M. Two frogs, dark in color, with red
streaks down the back. One in blue covered jar, one control.
June 18. 10 A.M. Frog much lighter than the control.
The red streak along the back now sandy colored, however,
.•<till with the reddish tint. Control color unchanged. Noticeably
darker than the one in blue.
The frogs reversed.
June 19. Lighter, the red streak sandy colored, same width as
before. Sides light grayish green. Whole tone lighter than the
one in the control.
The frogs reversed.
June 20. Lighter in tone, more greenish tinge. The red
.-treak now sandy, slightly greenish also.
I observed some pigment cells under the microscopes. The
melanophores (black) were the most noticeable, although on close
observation yellow and bluish grey pigment cells could be seen.
I stimulated the piece of skin with ice: in some cases the black
cells seemed to expand and in others this could not be seen. Some
such action, however, would be necessary to cause the darkening in
color brought about by cold.
The stimulations with heat were somewhat less definite than
with cold ; however twice the contraction of the melanophores, due
to a heat stimulus, was observed.
Left the two dead frogs for six hours. When I observed them
again they were both remarkably lighter than they had l:)een when
they were killed.
I took a bit of their skin and observed it again. One portion
was much lighter and had several isolated melanophores. I cooled
this piece of skin with ice, then stimulated it with hot water. The
]iigment cells seemed to expand.
Conclusions :
1. The tree frog changes its color in response to heat, cold
and changes in the color of its environment.
.^0 Journal of Entomology and Zoology
2. The frog docs not actually change color so much as it gets
lighter or darker in response to stimuli. There seems to be, how-
ever, some actual change in color.
3. The colors, blue, green and yellow cause the frog to get
lighter in color. The results with red were so irregular as to sug-
gest that the change might be due to some other agent than the color
environment.
4. When there is a reddish color present, i. e., red streak, the
red environment intensifies this coloration. When, however, there
is no red color present the red environment does not develop it.
5. This same phenomena is true of green coloration. Thus a
red frog does not seem to be able to change into a green one. nor a
green frog into a red one.
6. The changes in coloration or intensity seem to be due to
the expansion of the pigment cells.
Experiments with Rana sp.
I. May 26. Light. I left the frog in the light (sunlight,
although not direct) for one hour. At the end of this time it was
remarkably lighter than the one in the dark room.
II. May 26 Dark. Frog like the one in light. I left it in
the dark room for one hour. At the end of this time it was much
darker than the one in the light.
Reversed frogs.
Left two hours. At the end of this time the two frogs were
the same color again.
May 28. Repeated the first step of the light and dark experi-
ment with the same results.
May 29. Placed one frog on a white reflecting surface but not
in the sunlight. In one hour very little change.
At the end of the hour, placed the frog in the sunlight, still
on a white, reflecting surface. Remained there for one hour. At
the end of this time it was very much lighter than the one in the
semi-darkness.
Sunlight then has more effect than diffu.sed light, or perhajis
the difference is caused by the difference in temperature.
III. May 29. Placed one frog in a rather dark but not abso-
lutely dai'k place, used rather as a control than as an experiment.
Apparently it did not change color.
Left it for another hour. The supposition was that it did not
change color in the second hour, since the first hour had no effect.
However at the end of the hour it was much darker than the
one in the sunlight.
PomDii.i C()llc>zc, Clanmont, California 31
IV. May 27. Heat and cold. Placed a frog in water of 30
C, left it for an hour and one-half.
At the end of thi.s time it was decidedly lighter.
Placed a frog in water of 3 C, left it for an hour and one-half.
At the end of this time it was decidedly darker.
There was a great deal of difference in the color of the two
frogs at the end of the experiment.
V. May 28. Frog in water 30 C. Left one hour. Much
lighter than one in cold.
Frog in water 3 C. Left one hour. Much darker.
Reversed the frogs at 2:24 o'clock.
At 2:45, the two frogs had reached the same color.
VL June 1. Frog in water of 30 C, left one hour. Much
lighter.
Frog in water 3 C, left one hour. Much darker.
VIL May 26. Acid. Placed one frog in a weak acid (HCL)
solution. Left for several hours. There seemed to be no change
in color — possibly a little lighter than the control.
Control. Placed one frog in water, otherwise its environment
was the same as the one in acid.
No change in color.
VIIL Alkali. Placed one frog in a weak alkali (NaOH).
Left it for several hours. There seemed to be no change in color —
possibly slightly darker than the control.
EXPERIMENTS WITH A LOCAL FROG
IX. Rami draytonii
May 27.
X. Cold 3 C. Found a frog among the other.s identical in
color.
Left in cold for one hour. Darker at the end of this time in
comparison with the control.
XL May 29. Light. Placed Ra)ia draijtO)iii in sunlight for
an hour and one-half. At the end of this time it was very much
lighter.
Control. Kept the control in semi-darkness. Did not change
color.
Conclusions :
L These frogs change color under certain conditions of heat,
cold, light, dark, or excitement. Acids and alkalis have little if
any effect.
2. a. Heat and light cause the frog to lighten in color.
There is evidence that heat is the true agent, and light only as it is
associated with heat.
b. Cold and dark cause the frog to darken in color.
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VOLUME FIFTEEN NUMBER THREE
JOURNAL
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ENTOMOLOGY
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ZOOLOGY
SEPTEMBER, 1923
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT o/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
Notes on the Lepidoptera of Southern California No. 1
Donald C. Meadows 33
A List of Coleopetra Collected on the Beach During the
Summer op 1921 at Lacuna Beach— Clifford T. Dodds . . 35
Some Common Chinese Mollusca— Art/iwr S. Campbell 37
The Nervous System and Sense Organs, XIV— TF. A.
Hilton 43
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Notes on the Lepidoptera of Southern
California. No. 1
Donald C. Meadows
Two days during the second week of April 1922 were spent
collecting Lepidoptera at Corn Spring, Chuckawalla mountains,
Riverside county, California. The Chuckawallas are typical Colo-
rado desert mountains, being low and rough, and having the vege-
tation for the most part confined to sandy washes. Corn Spring
lies on the north side of the range in a canyon of the same name.
It is a small palm covered oasis having many introduced plants
as it is the home of an old prospector, who has a house and garden
at the spring. The elevation is approximately 1500 feet.
Fourteen species of butterflies were collected and three ob-
served. The nomenclature used is that of Barnes and McDun-
nough's Check List.
1. Pier is protodice, form oernalis — Edw. Three males and
two females taken. Fairly common around spring.
2. Nathalis iole — Bdv. Five males collected. Found spar-
ingly flying over bare, windswept desert mosaic. One specimen
taken near mouth of Corn Spring canyon far from any vegetation.
3. Eurymus eurytheme. form kcewaydin — Edw. Two males
and two females taken. Common near spring.
4. Danais archipinis — Fabr. One specimen seen at spring.
5. Danais berenice — var. strigosa — Bates. One specimen
seen with the above flying among the palms at Corn Spring.
6. Melitaea Neumoegeni — Skin — Wright. Fourteen males
and five females of this interesting species were taken. Probably
the most common butterfly of that locality.
7. Chlosync calif ornica — Wright. Nine males and five fe-
males taken in a small canyon about two miles above the spring.
These butterflies seemed to be very local in their distribution, one
small canvon being the only place that they were found. Types
figured by Wright from specimens taken in Colorado Desert, South-
eastern California. The Chuckawallas are at the northern edge
of the type locality.
.H Journal of Entom<)log>- and Zoology
8. Vanessa cardui — Linn. A few specimens seen flying in
Corn Spring canyon.
9. Apodemia mormo — Feld. One female taken.
10. Apodemia virgulti — Behr. One male taken flying with
the above. Contrary to expectations these two species were not as
common as in other parts of the desert.
11. Cah'philis netnisis — Edw. One male and two females
taken in canyon about two miles above spring.
12. Atlides halesus — Cramer. One female taken. Only one
other seen flying around a species of mistletoe.
13. Brephidium exilis — Bdv. Few Lycaenidae were found.
Two B. exilis were taken flying over grass growing near spring.
14. Hemiargws hanno — Stoll. Two males taken near spring.
This is a Mexican butterfly and only occasionally reported from
California.
15. Hemiargu.s isola — Reak. A male taken in canyon above
spring.
16. Pyrgus tessellata — Scud. A common butterfly through-
out desert. Very common around Corn Spring.
17. Thanaos clitus — Edw. Another common species in vicin-
ity of spring. A very fast flyer and difficult to catch. Six speci-
mens taken.
In all sixty seven specimens were taken near the spring.
A List of Coleoptera Collected on the
Beach During the Summer of 1921 at
Laguna Beach, California
CLIFFORD T. 13013DS
Determined by Dr. E. C. Van Dyke of the University of California.
CICINDELIDAE
Cicindela trifaaciata Fab. var. sig»ioidfa Lee.
CARABIDAE
Dyschirius marinuji (Lee.)
Bemidion ephippigerum (Lee.)
Bembidion indistinctum Dej.
*Bembidion cautum (Lee.)
Platynus californicus (Dej.)
HYDROPHILIDAE
Ochthebius hitcrniptus Lee.
tCercyon finibriatu-'i Mann.
STAPHYLINIDAE
Blediiis fenyesi Bnhr.
Cafius canescens Makl.
tCafius lithocharinns Lee.
tCafius luteipennis Horn.
Thinopinus pictus Lee.
XHadrates crassus (Mann.)
Baryodma sulcicollis Mann.
HISTERIDAE
tAcritiis maritimus Lee.
Saprinus scisstts Lee.
Saprinus bigevimeus Lee.
tSaprinus sulci frons Mann.
*This species is not recorded as being as far south as California
in Leng's (Catalogue of The Coleoptei-a.
36 Journal <if Knt<)ni<ilot;\ ami Zoolof^y
MELYRIDAE
Endeodes basal is (Lee.)
ANTHICIDAE
Anthiciws californicus Laf.
Anthicus maritimus Lee.
DERMESTIDAE
tDermestcs marmorattis Say.
TENEBRIONIDAE
Eulabis obscura (Lee.)
Phaleria limbata (Horn)
CHRYSOMELIDAE
Diacluis auratuj^ (Fab.)
CURCULIONIDAK
Phycocoetes testaccus Lee.
JThe names thus elieeked are rceorded by Lea Myers. Coleop-
tera From The Claremont-Laguna Region. Jour. Ent. and Zool.
1918. Vol. X. No. 3. pp. 4:3-53.
Some Common Chinese Mollusca
Arthur S. Campbell
During the last year I had the opportunity to collect and ex-
amine a number of the commoner littoral and freshwater shell-bear-
ing Mollusca occuring near Canton and at Chung Chow, Hongkong
territory. The shells enumerated include only a fair sample of
what might be obtained after longer search under more favorable
conditions.
It is interesting to note the alliance of this fauna with that
of the islands of the Pacific and with that of the California coast.
A number of species occur here that are found on the opposite
shore but there is a very complex admixture of the more definitely
warm-water forms and with some species of endemic origin. The
observations of Ralph Arnold (Palae. San Pedro, Calif., Acad. Sc.
O,")) concerning the tertiary shells of San Pedro and Japan shows
us the aflSnities at once of the living shell-bearing mollusca of
these two regions and likewise adds to our observations concerning
the relationship between the whole Pacific molluscan complex.
The molluscan fauna of South China appears to be paleotropical
considered in its broadest aspect.
All shells were determined by Dr. H. A. Pilsbury of the Phila-
delphia Academy. In all there are one hundred and thirty-seven
species represented in this collection.
(Contribution from the Zoological laboratory and Museum of
the Biological Survey of South China, of Canton Christian Col-
lege).
Gastropoda
Bullidae
Bulla ampulla L.
Acmaeidae
Helcioniscus eucosmia Pils.
H. toreuma Rve.
Haliotidae
Haliotus diversicolor Rve.
Turbinidae
Turbo coronatus var. granulatus Gmel.
T. intercostalis Pils.
T. japonicus Rve.
Neritidae
Nerita lineata Gmel.
N. undata L.
N. crepidularia Lam.
N. albicilla L.
38 Journal of Entomology and Zoology
Solariidae
Architectonica perspectiva L.
Littorinidae
Littorina irrorata Say.
L. palliata Say.
Viviparidae
Viviparus rossgeri V. Mlldff.
V. ciliata Rve.
V. orientalis Lea.
V. chinensis Gray.
V. aeruginosus Rve.
Cerithiidae
Cerithium morus Brug.
Clava sinen-sis Gmel.
Melaniidae
Melania ebeniiia Brot.
Stombidae
Strombus pugilis var. alatu.s Gmel.
S. canarium L.
S. succinct us L.
S. bittatus L.
Turritidae
Turris desbayesii Doumet.
Cassididae
Cassis japonica Rve.
C. inflata Shaw.
C. strigata Gmel.
Doliidae
Tonnia allium (Soub.) Dillon.
Pyrula dussumieri Val.
P. ficus L.
Cypraeidae
Cypraea arabica L.
C. carneola L.
C. errones L.
C. moneta L.
C. erosa L.
C. helvola L.
Muricidae
Murex torrefactus Sowb.
M. adustus Lam.
M. fulvesceiis Sby.
M. tribulus L.
Rapana bulbosa Sol.
Cymalium (Turrotriton) pfeifferiana Rve,
Gyrineum tubercuiata Br.
Pomona College, Clareiiiont, California 39
Thaisidae
Thais luteostoma Dillon.
T. lapillus L.
Nyctilochidae
Bursa rana L.
Distortrix reticulata Link.
Columbellidae
Columbella versicolor Sby.
Buccinidae
Buccinum undatum L.
Eburna lutosa Lamb.
E. areolata Lamb.
Alectrion obsoleta Soby.
Buscyon perversa L.
B. (Sycotypus) canaliculata Say.
Trochidae
Monodonta labrio L.
Tegula rusticum Gmel.
T. nigerrima Gmel.
T. argyrostoma Gmel.
Astraea undosa Wood.
Volutidae
Mitra aurnita Desh.
Olividae
Olivella sayana Rav.
0. (Callianax) biplicata Sby.
0. scripta Lam.
Gonidae
Gonus suturatus Rve.
Gonus carinalis Hw.
Gonus sulcatus Hw.
Turritellidae
Turritella Itacillum Kiener
Helicidae
Eulota similaris Fer.
Polygyra albolabris Say.
Gamaena cicatricosa Mull.
Gyclophoridae
Gyclphorus elegans Mldff.
Pyramidellidae
Pyramidula alternata Say.
Naticidae
Natica (Polinices) mamilla L.
N. P. melanostoma Gmel.
N. P. didyma Bolton.
Sinum neritoideus L.
40 Journal ot Entdinology and Zoology
Auriculidae
Melanpus liiteus Ciioy.
Scalidae
Epitonium lamellosa Lam.
Siphonaridae
Siphonaria japonica Don.
S. cornuta Gld.
S. sirius Pils.
Pelecypoda
Arcidae
Area (Scapharca) campechien.sis Gmel.
A. decussata Sby.
A. obtusa Rve.
A. granosa L.
A. (Brahatia) fusca Brug.
Parallele])ipedum torta St. March.
Mytilidae
Mytiiu.s .smaragdinus Ch.
M. californicus Conrad.
M. edulis L.
Modiolus fortunei Dkr.
Septifer virgatus Wiegen.
Pinnidae
Pinna incurva Gmel.
Atrina tuberculosa Sby.
Pernidae
Malleus albus Lam.
Ostreidae
Ostrea lakerousi Lamb.
0. cristata Born.
Pectinidae
Pecten pyxidalus Boru.
P. circularis Sby.
P. circularis var. aequisulcatus Cpr.
P. gibbus var. irradians Lam.
Amusium pleuronectes L.
Spondylidae
Spondylus cruentata Lisch.
S. imperialis Chemi.
S. sinensis Sby.
Unionidae
Anodonta woodiana Lea.
Pomona College, Clarcmont, California 41
Venei-idae
Tapes variegata Handley.
T. tristis Lam.
T. phillippinarum A and R.
T. phenax Pils.
Tivela stultorum Maue.
Gat'arium divaricatum Gmel.
Venus (Chione) cancellata L.
V. C. thiara Dillw.
Mactridae
Mactra (Spirilla) solidissima Dillw
Cardiidae
Cardium robustum Sol.
C. rugasum Sby.
C. sinensis Sby.
Chamidae
Chama rubea Rve.
Myidae
Corbula erythrodon Lamb.
Solenidae
Solen grandis Dkr.
Tellinidae
Tellina alternata Say.
Metis balaustina L.
Paphia striata Lam.
Caecella cumingi Desh.
Cyrenidae
Corbicula fuscata Lam.
C. fluminea Mull.
Ptericolidae
Ptericola pholadiformis Lamb.
Anomiidae
Anomia sir iplex D'Orb.
XIV. Echinodermata
ASTEROIDEA
The nervous system of the starfish is about the same in all
forms which have been studied. Only minor unimportant differ-
ences can be recognized and some of these may be due to the differ-
ent conditions under which the observations were made or the
different methods employed.
Along the radial and circumoral ambulacral vessels on the
oral side is a median thickening of the surface epithelium. This
is the chief part of the nervous system, that is the superficial
radial and circumoral system. Separated from these portions by
connective substance there are in each arm on each side of the
middle line the deep radial bands while within the nerve ring
about the mouth there are two deep circumoral bands continuous
with the two in each arm.
From the superficial nervous system fibers may be traced
directly to the surface layers of the tube-feet. From the inside
nerve rings, fibers follow the ambulacral system. The superficial
system is merely a thickening of the epidermis in certain regions
while the deep system is a thickening of the surface of the ambu-
lacral system. Nerve strands from the circumoral rings, proba-
bly from the deep rings, run to the stomach and other viscera.
In addition to the parts of the nervous system just described
there is a rather diffuse network of fibers and probably cells, found
in the body-wall outside of the muscles. This last has been called
the coelomic.
Sense cells and perhaps something of a nerve plexus seem to
be present below the epidermis all over the aboral and lateral parts
of the starfish. Just what relationship all these parts of the ner-
vous system bear to each other or how they may be distinguished
from each other, has never been made entirely clear.
Almost any portion of the body seems to be sensitive to touch
and there may be other sensations without special organs for their
perception. At the tip Of each arm a little tentacle or papilla
marks the end of the radial canal and the superficial nerve cord.
This little organ has a special epithelium and may be a special
organ of touch but Eimer, 1880, considers it as an organ of taste.
The eye-spot is the most marked sense organ of the starfish.
Each arm has. very near the termination of the radial nerve at the
tip of the arm, a bright red spot of pigment. A little closer exam-
ination of one of these spots shows it to be composed of a number
of distinct regions of color. In section these little areas are seen
to be little follicles lined with epithelial cells. The cells which line
the follicles are spoken of as the visual cells. These are clear at
their inner margins but pigmented farther down. Their inner
processes come into relation with the nerve strands at the bases of
44
Jinirnal of Entomology and Zoology
the eye-spots. Between and surrounding the visual cells are
numerous bipolar, elongate supi)ortive cells which stain strongly
with connective tissue stains. In some cases the eye areas are
not in the form of follicles as Pfefifer, 1901, has shown in a species
of AstropectcH. In those eye areas which appear as follicles a
lens has been described and figured by Pfeffer and others but I am
inclined to the interpretation of Cuenot, 1887, who believed that
no lens is present. In fact, in some sections which I have seen
there was no sign of even a membrane over the mouth of the
follicles.
In the superficial system many long supportive cells help to
make up the bulk of the nerve cord. These stain deeply with
usual stains and at their inner ends are more or less intertwined.
Fig.
A. DiaRi-am of a Starfish cut so as to expose internal as well as
e.\ternal parts of the nervous system. In the center the deep nerve
ring is shown liy a dark curved line, the surface nerve ring by
a thicker line. These parts are continued into the arm cut lonpi-
tudinally on the rijrht. Nerves to the tube-feet are shown. The
superficial nerve ple.xus and internal nerves are indicated. B.
Cross section of the radial nerve of starfish, superficial and deep
parts shown. C. Nerve cells and supportive cells from the central
nervous system. D. Section throujrh one of the pedicellariae of sea-
urchin showing distribution of nerves, after llamann. K. Section
through "taste knob" of eea-urchin. Hamann.
Pcim iiia C^dlcjic. Clarcinoiit, California 45
In the past I have been inclined to consider these as in part at
least with conductive function, but I am sure the true nerve cells
are sometimes bipolar, possibly in some cases multipolar with
fibers running longitudinally and laterally in the nerve strands.
The true nervous elements are more delicate, their fibers or fibrils
cross each other at various angles but bear no other obvious
relations to each other.
Among the earliest works on the nervous system and sense
organs of starfish is that of Hacckel in 1859. In 1860, Wilson has
a remarkably clear and accurate paper on the nervous system of
the starfish. Another early paper was by Owsiannikow in 1871.
Teuscher in 1856, figures the nervous system but not in much detail.
Ludwig, 1878, has his figure of the nervous system in section often
copied. Hamann, 1883-5, shows more of the structure of the
nervous system and gives a good idea of the structure of the eye.
Cuenot. 1887, gives a clearer idea of eye structure but not much
more about the detail of the nervous system. Jickeli, 1888, recog-
nizes four chief parts of the nervous system of starfish: (1) The
ambulacral, (2) the sub-epidermal body plexus, (3) the deep
nerves, (4) the intestinal nervous system. Pfeffer, 1901, studies
the eyes particularly and distinguishes clearly between support-
ing cells and nerve cells. More recent papers of Pietschmann,
1906, and especially of Meyer, 1906, show details in the nervous
system. The last author distinguishes clearly between supportive
cells and nerve cells in the nervous system. He finds the suportive
cells uni- or bipolar and usually running from the ventral to the
dorsal side of the nerve bands. The nerve cells are bipolar or
multipolar with fine branches.
Romanes, 1885, found besides strong negative reactions against
injurious stimuli, positive reactions of a chemical nature which
he considered due to the sense of smell. This sense depended
somewhat on the physiological condition of the animal, chiefly
upon its degree of hunger. A starfish, kept several days without
food, immediately crawled near some presented. If a small bit of
food be withdrawn as the animal approaches, the starfifish could
be led about in any direction. By severing various parts of the
rays, Romanes found that this so-called olfactory sense was equally
distributed throughout the length of the body and by varnishing
the upper surface he found that the reactions were unaff'ected.
Also by placing a bit of food on the alioral surface no reaction
occurred. Preyer, 1886, found great difi'erences in individuals
when stimulated with food.
Starfish are positively phototropic but largely lose this ten-
dency if the eye-spots are removed. Romanes found the sensi-
tiveness so great that starfish discriminated between ordinary pine
boards covering the tank and the same boards painted black.
Romanes Preyer, Jennings and others have studied the righting
46 Journal of Entomology and Zoology
reactions of starfish in considerable detail. In general the star-
fish rights itself by twisting about two or three of if>- rays until
the suckers on the ventral side have a firm hold of the supporting
surface and by controlling the twisting movement the l-oily is
turned over. In this it is necessary that all five arms do not make
the attempt at once to bring the animal into a ventral position. If
five or four arms should work at once the animal could not turn
over. There must be some coordination between the arms as is
seen when the circum-oral nerve is cut. In this case the coopera-
tion of the arms ceases. A single arm removed from the rest can
right itself. These experiments seem to show that the central
nerve ring acts merely as a conductor of impulses. The ventral
side of the starfish seems to be positively stereotropic.
If one arm of a starfish is stimulated the animal moves away
in a direction opposite to the stimulated arm. This looks like
intelligence, but when one arm is stimulated the tube-feet on this
arm draw in and according to the parallelogram of forces, a move-
ment away from the point of stimulation will take place. When
the starfish is stimulated as a whole the spines and pedicellariae
wave about and the jaws snap time and again. A separate exter-
nal stimulus is not necessary for each opening or closing of a pedi-
cellaria. Mechanical stimuli that are ."Strong enough always cause
them to attack. Very light mechanical shock often produces no
effect even if repeated. There are some responses to food rather
than defensive movements, a nutrient juice causes the pedicellariae
to advance and open. Pedicellariae are often opened for attack.
If another starfish brushes against it, even when one of the indi-
vidual's own rays cro.ss, the pedicellariae may be advanced.
If closed pedicellariae are stimulated they mu.st first be stimu-
lated to open i)efore they will attack. Any stimulus which cau.ses
the pedicellariae to rise will when repeated cause them to open.
Most stimuli which cause the pedicellariae to withdraw also cause
them to close. The larger pedicellariae are usually less inclined
to hold objects for a long time. Starfish seem to hold objects for
a longer time than sea urchins.
In starfi.sh the pedicellariae .seize and hold each other as well
as other objects. If a small bit of the body of a starfish, bearing a
single spine be cut from the rest, the pedicellariae seize any small
object which touches them. If the ventral nerve is cut or the
whole ventral side of the ray cut the pedicellariae continue to act,
but the cutting of the nerve acts as a stimulus. The transmission
of impulses seems to be by the nerve-nets over the body-wall.
Jennings has shown that the elevation of the groups of pedi-
cellariae or the rosettes to attack, is dependent upon the following:
1. Previous mechanical stimuli; 2. Preliminary chemical stimuli;
.?. Foregoing chemical stimuli; 4. Cutting the radial nerve leaves
the ro.settes in such a state that they attack more readily than
usual. 5. The rising of the rosettes in a central region leaves them
^^^
Fig. 28. Sense organs of Starfish. From Campbell.
1. Ventral and lateral views of eye-pad Pisaster capita tus, showing
general relationship to terminal tentacle. X9.
2. Ventral view of eye-pad of Ortliaster gnnolena. X9.
3. Ventral view of eye pad of P/sosfer oc/irocei(S. X9.
4. Ventral view of eye-pad of Asterina miniata. X9.
5. Ventral view of eye-pad of Lmckia colombiae. X9.
6. Ventral view of eye-pad of Asteropectin erinaceus. X9.
7. Ocellus from Orthaster gonolena to show general form. X350.
Drawn by camera lucida.
48 Journal of Entomology and Zoolog)'
8. Ocellus from Linckia colonibiac to show general features. X350.
Camera lucida.
9. Ocellus from Astcriiia mhuatn. X350. Camera lucida. General
view, note the clear central margin of pit.
10. Tactile organ from terminal tentacle of Linckia colombiae.
General view showing papillae and details. Camera lucida.
X350.
11. Single sensory cell from Linckia colombiae. Very greatly
magnified.
12. Sensory cells from Asterias riihens showing pigrment. Re-
produced from Cuenot. Osmic acid. Greatly magnified.
13. General view of eye-pad of Asteropectin eriuaceits. X350.
Camera lucida.
14. Simple ocellus in an Asterias. Supportive cells dark. Sensory
cells lighter. Reproduced from Pfeffer. Diagramatic.
15. A more complex ocellus from Asteropectin m.ultcri. Note the
lens, other features as above. From PfefFer. Diagramatic.
after subsidiance in such a state that they react more readily to
stimuli in a distant part of the body than the rosettes near the new
stimulus; 6. There are differences in the characteristics of indi-
viduals.
The opening of the pedicellariae depends upon :
1. Homogeneous preparatory stimuli
(a) Sometimes there is no response to the first stimulus.
(b) Sometimes the first .stimulus causes retraction and
closing while later ones cause e.xtension and opening.
(c) Sometimes with large pedicellariae the first stimulus
causes momentary opening, the next two or three have no visible
effect, the next pronounced opening.
2. Chemical stimuli of a certain character cause the pedi-
cellariae to open later and more readily under mechanical stimuli.
3. Chemical stimuli of a certain character cause later refusal
to open under usual mechanical stimulation.
4. Holding some object causes the pedicellariae after release
to refuse to open under other stimuli.
5. After closing the pedicellariae often open and close again
spontaneously, "snapping." The foregoing action furnishes the
condition for the succeeding one.
In many cases the tube-feet are compelled to do much feeling
about before they find the object seized by the pedicellariae. In
oxjiloring movements two or three rays are raised from the others
and swung about in the water; the other rays creep about. The
tip of the arm as well as the other parts of the arm are employed
in these feeling motions.
The relative intensity of illumination on different parts of the
body of the starfish may and at times does determine the direction
of movement without regard to the direction of the rays of light.
The ventral portion of the surface of the .starfish is protected by
Pomona College, Claremdiit, California 49
movements more than the tips of the arms. After it has been at
rest for a time however the eye-spots are usually so placed as to
be protected from the light. The starfish in each case (Jennings)
moves towards that part of the body that is least illuminated.
There are a number of ways in which starfish right them-
selves according to Jennings :
1. The simplest method. Two adjacent arms twist their tips
with ventral faces inwards.
2. Two arms, the ventral faces not inwards but facing in the
same direction.
3. Three adjacent rays attack and usually turn by twisting
the outward rays.
4. Four rays take hold, two to right, two to left. Fifth ray
helped up, and swings over.
5. All rays attack release later of certain rays.
6. One ray twists and rights the whole.
7. Righting without attaching tube feet of any of the rays.
Raises disc strands on tips of arms then topples over.
If a starfish begins a reaction in a certain way it usually con-
tinues in the same way even in spite of opposing conditions.
When the starfish gets started it continues in the same way. The
variability of form in starfish that are righting themselves is
very great. No species rights itself in one way alone. When cer-
tain tube-feet are prevented from acting in righting movements
the others carry on the movements. In righting if one method
does not help another is used. ,
Habit Formation
Preyer, 1886, Jennings, 1907, have brought further information
as the results of experiments to test habit formations in starfish.
By perventing certain rays to act in the righting reactions in star-
fish Jennings showed that he could establish temporary habits
and the slower formation of more lasting habits. The many
factors which determine the righting reactions have not a constant
tendency to make starfish turn on one given pair of rays. On the
contrary, they must sometimes act in one way, sometimes in an-
other. Therefore nq very fixed habits are formed under normal
conditions.
In the righting reactions the impulse tends towards the ac-
complishment of the general turning of the starfish as a whole and
given parts sacrifice their own direction or even prevent it in the
general result.
We cannot assume single specific external stimuli as the deter-
mining factors for each separate movement, yet in some way the
relation of the organism to its environment has set in operation a
uniform action of which separate movements are parts.
50 Journal of Entomology and Zoology
ECHINOIDEA
The nervous system of sea-urchins may be compared with that
of starfish more ea-sily than with that of other forms.
The nerves corresponding to the superficial radial and circum-
oral nerves are more deeply placed than in starfish and as in star-
fish are the most obvious parts of the nervous system. An epi-
neural space or tube on the outer side of the nervous band forms
the so-called "epineural cavity" or nerve tube, as interpreted by
Phouho. '87, and others. The radial and circum-oral sinus follows
the nervous system on the inside.
The superficial radial system follows down the inside of the
shell in the center of the ambulacral area and these five strands
join with the circum-oral ring about the mouth opening.
From the nerve ring between the junctions of the five radial
nerves are branches to the intestine which go to make up the intes-
tinal plexus. Nerves run out laterally from the radial nerves to
the tube-feet and also to the surface, to the bases of the spines and
to the ganglia at the bases of the spines. The radial nerves end
in the terminal tentacles through holes in the shell about the anal
region. It is by way of these openings, according to Phouho, that
the radial nerves contribute to the superficial nerve plexus just
outside the test of the sea-urchin. The deep radial nervous system
is but poorly represented, so little of it is present clo.sely applied
to the superficial radial and circum-oral that it can hardly be recog-
nized apart from it.
According to some, a pentagonal area of aboral nerves sur-
rounds the anus and communicates' with the genital organs and
with the external superficial .system by means of fine fil)ers from
the radial nerves near their termination in the terminal tentacle.
It is quite prcjjable that the superficial system communicates with
that of the shell at the aboral end not only through the so-called
ocular openings but also through the genital openings in the shell.
The surface of the body, the spines and the tube-feet, are all
organs of the tactile sense at least.
The so-called eye-spots at the terminal tentacle in the five ocu-
lar plates contains pigment and may have some sensitiveness to
light, but it is not like the eye-spots of starfish and may indeed not
be in any sense an eye-spot.
The chief parts of the system such as the radial and circum-
oral nerve bands are composed of about the same parts as in the
starfish. In smaller and perhaps younger specimens the outer
nuclear layer is thicker in proportion. Nerve cells are bi- and
multipolar. In some cases at least multipolar cells are found well
within the fibrous area of the strand. Many of the outer cells are
probably as in other echinoideans supportive in function. The
radial bands are thicker at the oral region and become somewhat
Pomona CoUi-^e, Clarcniont, California
51
smaller at the I'egion of the terminal tentacle in the ocular plate.
This might suggest something as to the nerve tracts or bundles of
fibers and gives an indication at least that fibers may convey im-
pulses at different distances such as in the central nervous system
of vertebrates.
The deep radial and circum-oral strands of sea-urchins are
poorly shown in section. Only a few cells scattered along the
inner margin of the fibrous region give an indication of this poorly
developed system.
In the sand-dollar, Dendraster excentricus some variation in
form is suggestive of value in comparison with other forms.
The righting reactions in sea-urchins are carried out with
greater difficulty than in starfish and only the fresher or more vig-
orous individuals are capable of the reaction.
Fig. 29. Nervous system of Sea-urchin. A. Diagram of nervous system of
sea-urchin showing in various ways the superficial and deep nerv-
ous system by having the superficial system cut away on part of
the two radial nerves at the left. Branches to the tube-feet shown
in the central of the three ambulacral areas. Nerves to the bases
of the spines show on the right. Superficial nerve plexus show in
the center. B. Diagram of the nervous system from the aboral
pole, showing the nerve connections at the genital openings and the
ends of the radial nerves at the five ocular plates. C. Diagram of
cross section of nervous system having branches to a spine and
a tube-foot after Delage and Herouard.
Although the eye-spots of sea-urchins are not well developed
they seem to avoid' light and seek darker corners and sheltered
places. One form which has no eye-spots seems to avoid the light.
A sudden shadow falling on it causes it to direct its spine to the
^(t? cat
Fig. ao. Explanation or Fici'nES of Sand-doixak.
1. Diagram of one fifth of Aristotle's lantern of />(Hrfrnxf<'i- show-
ing three loops of the circumoral nerve ring, and parts of three
railial nerves, the central one partly hidden at its origin by the
lantern. The nerves are in black. X'.t.
2. Drawing of part of the first part of an oral railial nerve. Xil.
:{. Drawing of the lower end of an oral ra<lial nerve. X9.
Pomona College, Claremoiit, California 53
4. Drawing of the upper part of an aboral radial nerve. The
eye-spot reg-ion is up in the figure. X'J.
5. Camera lucida drawing of a part of an aboral nerve showing
position of cell areas. X70.
6. Drawing of a section of an oral radial nerve. X300.
7. Drawing of a section of circumoral nerve. X300.
8. Drawing of a section of aboral nerve. X300.
9. Nerve cell.s from central regions of a radial nerve. The ar-
rangement is as shown in the drawing, cells of various levels shown
as one layer. Some of the processes possibly relate nearby cells,
but most fibers run into the general fibrous mass. All fibres or
fibrils are small. There is some indication of tigroid substance in
some of the cells. X450.
10. Nerve cells from near a lateral branch from the radial band.
X450.
shaded area. Uexkull, 1897, was of the opinion that the sea-urchin
possessed a special set of nerve fibers concerned with photic
responses.
If a bit of the test with one or more spines be separated from
the rest of the animal, the spine or spines may be stimulated to
react much as before. In the sea-urchins there are several kinds
of motile organs. There are the jaw-like organs or pedicellariae,
borne on movable stalks ; there are the tube-feet and the long mov-
able spines. All these sets of organs are controlled by nerves, and
a nerve network connects all these motile organs. One general
network of nerves is within the shell and one without, and these are
connected with the five radial nerves and the circumoral nerve ring.
Each of these motile organs has a definite number of reaction or
responses and in these each group may act independently and each
organ may react as an independent individual. Each sea-urchin
then seems according to Uexkull to be made up of a colony of
almost independent structures yet all these are connected by the
nerve network and when one carries out a reaction others may
receive a stimulus to carry out its special activity.
The independence of these systems of spines, pedicellariae and
tube-feet, and the definite character of their reflexes has been
clearly expressed l)y Von Uexkull He considers the sea-urchins as
made up of a "republic of reflexes." Each reflex is of the same
value and is independent of the others except for the nerve-net
connections between the systems. This group of chiefly independ-
ent systems has nothing like a central unity controlling them as a
whole and it is only by the synchronous course of different reflexes
that a unified action is simulated. The action is not unified but
the movements are ordered. Separate reflexes are so constituted
and so combined that the simultaneous but independent course of
reflexes in response to outer stimulus produces a definite general
action similar to the condition in animals with a common center.
54 Ji(iirii:il lit Entomology ;ind ZiK)lo^y
The pedicellariae of sea-urchins refuse to seize or hold each
other or parts of the bodies or others of the same species. Von
Uexkull believes this is due to a presence of a substance "auto-
dermin" which is in the skin. In sea-urchin pericellariae have the
power of indei)endent reactions. Each when isolated on a piece of
shell may behave as when on the body of the animal. The stimulus
from the pedicellariae need not pass through the radial nerves for
if the nerve is cut the reactions are as before.
Ophiuroidea
The nervous system of serpent-stars is more complicated than
that of starfish. The radial and circum-oral nerves are shut off
from the surface of the body and inclosed in a small cavity. The
more superficial radial and circum-oral nerves are well developed
and from the radial nerves fibers run out to the spines of the legs
and the tube-feet. These last are provided with ganglia at their
bases and with nerve strands running their length. The nerves
which run to the spines also have ganglionic thickenings upon
them at the bases of the spines. From the ganglia at the bases of
the tube-feet delicate strands run out to one epineural ganglion
for each tube-foot. On the ventral side of the disc on each side of
radial nerves lateral nerves run out to near the margin of the disc
communicating with the radial nerves and also connected with the
superficial nerve jjlexus on the lower side of the disc. There are
then in this way two lateral nerves from each arm base, and each
one of these sends out an inter-radial nerve. Nerves from the
circum-oral ring run to the teeth, five pairs in all.
The deeper radial and circum-oral nerves are closely applied
to the more superficial nerves and appear much like parts of them,
being represented by groups of dorsally placed cells. These deeper
nerves are two for each arm. The circum-oral ring about the
mouth sends out pairs of nerves to the muscles of the arm.
A system of so-called genital nerves is found quite distinct
and independent from the other systems. More or less isolated
loops of fibers are found in each area of the disc between the arms.
Ilamann, 1888. gives one of the best accounts of the nervous
system. Delage and Herouard also figure and describe the nerv-
ous system in this group. The first author describes and figures
nerves coming out laterally from the chief radial nerve to be
applied to the skin. These may be the cutaneous nerves of Delage
and Herouard.
Ilamann al.so shows strands from these to the tube-feet where
ganglia are located and from these ganglia are nerves with ganglia
running in to the center of the arm, and also nerves arching up
dorsally to end in small ganglia. These are very much in the posi-
tion as shown in the diagrams and figures.
Pomona College, Clarcmont. California 55
The chief radial nerves, as is well known, are concentrated
into ganglion-like swellings at the intervals between the vertebrae
and here from the dorsal nerve cord strands are sent into the mus-
cles of the arms.
Fig. 31. Nervous system of Sehpent-Stars. A. Diagram of the nervous
system of a serpent-star, a part of the disc and the bases of three
rays shown. In the upper right end ray the superficial nerve ring
and radial nerve are removed to show the deeper nerves. In the
others and on the disc other nerves are shown. On the disc the
superficial nerve-net is given. Out from the radial and circum-oral
nerve the chief branches to the tube feet, etc., are shown. B. Dia-
gram of the nervous system of one of the arms cut across showing
large cavity of an arm in deep shading and the lowest cavity
within radial nerve, with branches to spines and tube-feet. C.
Diagram of section of an arm after Hamann. D. Through the arm
at another level. E. Section through radial nei've. Hamann. F.
Section through sense papilla. Hamann.
56 Journal of Entomolojiy and Zoology
The parts of the nervous system are :
1. The superficial radial nerves. The chief branches: (a)
muscular nerves, (b) cutaneous nerves to tul)e-feet and to skin and
to spines. On each nerve to the tube-feet a ganglion is formed at
base of each tube-foot and strands run dorsally and centrally to the
intervertebral ganglia and ventrally to the two ventral ganglia or
epineural ganglia, (c) branches communicating with the lateral
nerves of the disc on each side of each radial nerve which in turn
have altogether 10 interradial nerves near the center of the disc
and branches to the superficial nerve plexus.
2. The superficial oral nerve is pentagonal in .shape and gives
off: (a) nerves to the stomach, (b) a pair of dental nerves.
3. The deep radial nerves give off nerves to the muscles of the
arms.
4. The deep oral ring gives off: (a) interradial superior
nerves, (b) interradial internal nerves.
5. The genital nerves, independent of the others. Five dif-
ferent nerves between each radial area in the disc.
There are no eyes. The skin has no cuticle except at certain
jioints and these are the only ones where sense cells are located.
The tube-feet and spines are sensitive to touch. The palps are
.sensitive to touch as well as parts of the general surface. The
extremity of the palps have sensory functions. The terminal
tentacle, it is thought, may be olfactory in function. The oral
palps have sensitive papillae well supplied with nerve cells.
The structure of the nervous .system is somewhat like star-
fish but the central cords are parallel with more evident nerve cells
and the strands seem to have a more complicated structure.
Hamann's work is perhaps the most valuable in this group.
Delage and Herouard make chief use of this in their work. Teus-
cher 1876, Land, '7G, Smith, '79, and Ludwig, '80, are the other
chief contributors who have considered the nervous .system.
NOV 17 1939
fc
^ji
VOLUME FIFTEEN NUMBER FOUR
JOURNAL
OF
ENTOMOLOGY
AND
ZOOLOGY
DECEMBER, 1923
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT 0/ ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
CONTENTS
Page
A New Salt Marsh Mealy Bug — Clifford T. Dodds 57
Notes on the Life History of Dinaparte Wrightii Horn
— Roy E. Campbell 61
Nervous System and Sense Organs XIV Cont.— W^. A.
Hilton 67
Entered Claremont, Cal.. PostOfflcc Oct 1. 1810, as second-class matter, under Act or Congress of
March i. 1879
Journal of Entomology and Zoology — Advertising Section
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A New Salt Marsh Mealy Bug
{Eriucoccus palustris n.sp.)
Clifford T. Dodds. University of California
Introduction. — While making a study of the insects of the salt
marshes and brackish waters of the San Francisco Bay region, I
chanced to find in considerable numbers, a mealy-bug on the salt-
marsh cord-grass {Spartina foliosa Trin.). It occurred on the
upper surface of the leaves and generally out of reach of the
ordinary high tides. The probable reason why this mealy-bug, as
well as the scale, Cliinaspis spartinae Comst„ occurs almost en-
tirely on the upper surface of the leaves is because of the fact that
during transpiration, water is given off from the lower surface
of the leaf, leaving after evaporation a considerable deposit of salt.
Type host and locality. — From Spartina foliosa Trin., at Al-
monte, Marin Co., California, November, 1921. Found wherever
the host is located about the shore of Richardson's Bay, an arm of
San Francisco Bav.
Fig. B. Leaf of host plant Spaiii}i(i foliosa Trin.; w, Cluunaspis spartinae
Comst., adult female; x, E. pahistri.'i, female before secreting sac;
y, sac that has been wet by the tide; z, normal sac.
Sac. — The natural sac is composed of fluffy white waxy fila-
ments (Fig. B, z), which after they have become wet by the tide,
and this is usually the case in nature, become a light ashy gray (1),
and have a more or less feltlike texture (Fig. B, y). thus offering
great protection, especially for the overwintering young. At the
posterior end of the sac there is an obscure opening, plugged with
wax filaments, where the young escape. The average length of
the sac is 4 mm. for the adult females and somewhat less for the
males.
Adult female. — type — (Fig. A) Body smooth; six cephalotho-
racic spines on the dorsum, the two median anterior ones being
larger than the other four, all straight, slightly expanded at the
base, tapering to a rather blunt apex; eight pairs of very small,
blunt, conical, dorso-lateral marginal spines on each side; the
58 Journal of Entomology and Zoology
posterior spine of the last pair of each marginal series slightly
larger than the others. On the ventral surface there are sparsely
scattered hairs, arranged segmentally ; four spiracles located pos-
terior to the coxae of the front and middle legs. Anal lobes not
chitinized, each with three small ventral and one large terminal
setae (Figs. C, D) and two dorsolateral spines on the inner sur-
face. These spines are slightly larger than the cephalothoracic
spines mentioned above, not expanded at the base and very blunt.
The last pair of the marginal spines are located dorso-laterally
near the basal end of the anal lobe (Fig. E). The terminal setae
of the anal lobes are about two and one half times as long as the
anal lobes themselves (Fig. C), while the eight setae of the anal
ring are less than the length of the anal lobes. Antennae medium
stout, six to eight segmented (Figs. G, H, I, J,), the normal long
Fip. A. t'rincocniK palustris n. sp., adult female cleared in caustic potash.
third divided into the third and fourth, and the normal ultimate
segment divided into the seventh and eighth. Apparently this di-
vision is not clo.sely related to the moults. Legs rather slender
(Fig. F), claws not toothed, digitules with flat apical enlargements.
Male. — Body 1 mm. long; folded wings projecting •'/|. mm. be-
yond end of abdomen.
Pomona College, Clarcmont, California
59
Eggs. — Average 60 to 70 eggs per female, 92 highest number
noted. Ellipsoidal, pale cadmium yellow (1) ; .5 mm. long, .25 mm.
wide.
Type and paratypes deposited in the California Academy of
Sciences, paratypes also deposited as follows: United States Na-
tional Museum, Washington, D. C; G. F. Ferris. Stanford Uni-
versity, Palo Alto, California; E. O. Essig, University of Cali-
fornia, Berkeley, California, and in my own collection.
Comparison. — This is a very distinctive species, the small
number of spines, their form, size and distribution separating it
quite widely from the known species of this state. The only
species that I have seen which at all resembles it is Eriococcus
inerTnis Gr., which is found on grass at Camberley, Surrey, Eng-
land.
Life history. — As a rule the females come to rest with the
cephalic end of the body uppermost on the erect leaves, where they
Fig. C. Anal lobe showing relative positions of dorsa-latoral spines to ven-
tral setae.
Fig. D. Venteral aspect of anal lobe.
Fig. E. Dorsal aspect of anal lobe.
Fig. F. Leg.
Figs. G, H, I, J, showing variation of antennae.
Note: (1) Nomenclature of Windsor and Newton's water colors as given
in the "Glossary of Entomology", Smith. Brooklyn Ent. Soc, Brooklyn, N. Y..
1906.
60 Journal of Entomology and Zoology
secrete the sac about themselves. The eggs are laved in the
bottom of the sac, being quite closely packed with wax -filaments.
As the female deposits the eggs her body is crowded forward, the
dead remains being found in the upper end of the sac. In some
instances in the laboratory, it was noted that undersized females
would secrete a sac, dejiosit a small number of eggs and die. Upon
dissection, it was found that there were often mature eggs in the
oviducts or partly developed eggs in the ovaries. The first laid
eggs in the bottom of the sac hatched several days in advance of
the others and thus the nymphs escaped before the later eggs
hatched.
Under laboratory conditions the first moult took place 17 days
after hatching and the second moult a week later. After the sec-
ond moult the individuals had lost the pale cadium yellow color and
became the violet gray of the adult. The day following the second
moult they secreted sacs, although they were very much smaller
than those which first made sacs under natural conditions. Nor-
mally they probably have five moults. The overwintering indi-
viduals found in nature were first instar nymphs which had not
left the sac.
Parasites. — One Hymenopterous insect was found, Pscudo-
coccobius clau.'iscni Timberlake (2) which parasitized a large per-
centage of the mealy-bugs. As many as si.\ of these parasites
were found in an adult female. They usually kill the female after
she has made the sac and before oviposition. They overwinter in
the sac as adults, emerging in the spring through circular holes
which they make.
Resistance to drowning. — Several experiments were made to
see if this insect was specially protected from the water. It was
found that submergence for three or four days had no ill effect on
an adult and that they could float on the surface of fresh water for
three weeks without dying. These results are of little significance,
however, as Mr. Floyd Wymore, in his B. S. thesis work at the
University of California, 1922, found that Pseudococcus gahani
Green, a terrestrial mealybug, not only could live under water but
laid eggs and otherwise lead (juite a normal life.
Acknowledfiements. — I am deeply indebted to Prof. G. F. Fer-
ris of Stanford University for numerous courtesies, especially for
determining this mealy-bug as a new species and for the permis-
sion to examine his collection of Coccidae. I am also indebted to
Mr. W. C. Matthews for photographing figure B, and to Prof. E. O.
Essig for suggestions and aid.
Note: (2) Mr. A. B. Gahan, F^ntomoloRical Assistant of the United States
National Museum, writes as follows: "The parasite appears to be Ptnitdo-
rorcnhiux cliiunsriii Timberlake. This species was described from a sinsle
male specimen bred from Eriitm. sp. [Lichtensioides Ckle.] * * ' at Riverside,
Calif. Your males fliffer very sliRhtly in the extent of yellowish color on
the face but I believe there is very little reason to doubt that they represent
this species with the type of which they have been compared."
Notes on the Life History of Dinapate
wrightii Horn. (Col.)"
By Roy E. Campbell, U. S. Bureau of Entomology, Alhambra, Cal.
In May, 1916, Mr. J. 0. Martin, of Pasadena, after consid-
erable tedious scouting, discovered a log of the Washington Palm
(Washingtonia plifera) in Palm Canyon, on the Northwestern
border of the Colorado Desert, which contained partially-grown
larvae of Dinapate wrightii. Mr. Martin could hear more larvae
at work in the log, and decided to mark it and wait until the follow-
ing spring for further action. In 1917 he returned to Palm Can-
yon and sawed out several pieces from the fallen log, carried them
down the canyon to his automobile (a feat which was discovered
a little later by the writer to be quite laborious) and transported
them to Pasadena.^
At the suggestion of Dr. F. H. Chittenden, the writer got into
communication with Mr. Martin and received directions to locate
the remaining 10 feet of the infested log. On May 19, 1917, the
writer went to Palm Springs and duplicated Mr. Martin's actions,
except that the scouting was unnecessary. The logs were placed
in a wire cage, in Alhambra, and closely watched. Mr. Martin's
efforts were rewarded by the appearance of the first beetle on
August 3, and emergences continued until thirty-one had appeared
by September 17. In the writer's cage 3 adults appeared in the
latter part of July, and 2 in August.
When the sections were sawed from the log, a few larvae were
disclosed, some practically full-grown, while others were quite
apparently immature, indicating the possibility of two broods.
Also after the emergence of the beetles in 1917, larvae could still
be heard at work within the log.
On April 15, 1918, one piece of the log which had been trans-
ported to Alhambra, was cut up and examined. Nine larvae were
found, four of which were full-grown, and the rest not over half-
grown. These larvae were sent to Dr. Chittenden. Also one dead
adult female, which had failed to make its way out of the log,
was uncovered. The emergences of adults for that year from the
remaining piece of log were as follows :
" Bull. Brooklyn Ent. Soc. Vol. XII No. 5, pp. 107-110, December, 1917.
*Since tils paper was piesenled for publication, an article by Dr. J. A. Comstock on "A
Giant Palm-Borins Beetle" appeared in the March, 1922. Bulletin of the Southern California
Academa of Sciences (Vol. XXI. PaTt I). Besides giving many of his observations, it reviews
the literature on this interesting beetle.
62 Journal of Entomology and Zoology
August 1, 1918—1 male.
August 2, 1918—1 male.
August 2, 1918—1 male.
August 8, 1918—1 male.
Sept. 2, 1918 — 1 female, elytra deformed.
Since there evidently was still another brood, or some larvae
were slower in developing, the remainder of the log was kept, and
three beetles emerged in 1919 as follows:
July 24, 1919 — 1 male, large fine specimen.
July 25, 1919 — 1 male, small specimen.
Aug. 25, 1919 — 1 female, average .specimen.
No further attention was paid to the log until April 1, 1920,
when out of mere curiosity, it was cut up. To the writer's great
surprise one live larva was found.
It did not appear to be quite full-grown, or at least was a little
undersized, and was soft and flabby. Although it was not ex-
pected that it could mature, a hole was bored in the end of a piece
of the log, near and parallel to the surface, the larva put in, the
hole corked up and the piece of wood placed upside down. The
larva soon began to bore into the wood.
On May 24 it was examined again. The larva had continued
boring into the wood, parallel with the bark, filling up the hole
behind it, and had turned around in the hole and was headed up-
ward. It finally worked a little to one side, and started upward
parallel with the other gallery. It was then transferred to another
piece of log. and put in a hole bored about 2 inches deep. During
the transfer, the photograph of the larva in the gallery shown in
Plate I, A, was taken.
The cork plug was removed frequently and the progress noted.
Not much eating was done after the above date, and on July 12
the writer was delighted to find that the pupa had formed. It
was creamy white, with dark eyes. By August 4 the legs, mouth-
parts and head were turning brown, and on August 8 the adult
formed. It was put back in the hole and the latter plugged up.
The beetle proceeded to the top of the gallery and ate its way up-
ward and outward. It emerged from the log on August 23, a
medium-sized female. The gallery eaten by the larva between the
time it was put in on May 24, and pupation on July 12, in which
pupation took place, is shown in Plate I. B — C, and the exit hole
eaten by the beetle at C — D. The walls of the gallery made by the
larva are much smoother than those made by the beetle as the latter
ate its wav out. An exterior view of the exit hole is shown in
Plate I, F."
A resume of the above indicates the following:
May, 1916, Palm log with immature larvae discovered in Palm
Canyon by Mr. Martin.
EXPLANATION OF PLATE I
A. Mature larva of Diiiapate wrightii in gallery just preparatory to
pupation. B — C. Parallel section of gallery eaten by larva between May 24
and July 12, in which pupation took place. C— D. Hole eaten by adult in order
to escape from log. E. External view of exit hole.
64 Journal of Entomology and Zoology
May, 1917, Log removed to Alhambra, California.
July and August, 1917, 5 adults emerged from log.
April, 1918, 4 full-grown and 5 partly-grown larvae observed
in one piece of log.
August and September, 1918, 4 adults emerged from re-
mainder of log.
July and August, 1919, 3 adults emerged.
April 1, 1920, one nearly full-grown larva found in log.
July 12, 1920. larva pupated.
August 8, 1920, adult formed.
August 23, 1920, adult emerged from log.
The partly-grown larvae observed by Mr. Martin in 1916 must
have been the ones to emerge in 1917 and 1918, indicating the life
cycle to be at least 2 or 3 years. However, they may have been
more than one year old in 1916. Mr. Mai'tin believes that the
small larvae observed in 1917 were from a brood deposited after
the log was discovered in May, 1916. However, it is apparent that
there was no deposition after the logs were taken in May, 1917,
and it seems probable that the latest deposition possible was from
beetles which emerged in the summer of 1916. If this is true,
then the life cycle of the beetle emerging in 1920 was practically
4 years. It is possible that deposition occurred prior to 1916,
which would make the life cycle 5 years or more. Beetles emerg-
ing in the other years must have been from 1 to 3 years old at the
time the log was discovered. If this is so, it would make 4 sep-
arate broods, which seems improbable. It is the writer's opinion
that there probably were two broods, and that the life cycle of
Dinapate ivrightii may vary from 3 to 5 years. It is certain that
the period can be four years. The quantity and quality of food
accessible to each individual larva no doul)t had much to do with
the rate of development, but probably other factors enter in also.
If the log contained only one brood, then the variation in the length
of life would be still greater.
It is interesting to note that when Mr. H. G. Hubbard visited
Palm Canyon in February, 1897, he observed that "all larvae were
thoroughly dormant and very flaccid. There are no young, and
evidently all are of the same age and nearly or quite adult. I
feel sure that they are more than one year old, and probably more
than 2 years old, but no doubt they would have issued in July or
August of this year."-
Specimens sent to Washington by Mr. Hubbard did emerge
in August. His belief that the life cycle would be at least 3 years
is demonstrated by the writer's experience.
' Ent. News, Vol. X, No. 4, pp. 228-230. 1899.
Pomona College, Clarcmont, California 65
Mr. Richard T. Garnett visited Palm Canyon on May 21 and
22, 1917, and after extended search, found an infested log, from
which he took 133 adults, 28 pupae and 17 larvae. One fresh exit
hole was observed. This and other observations indicate that the
period of emergence of the beetles extends from the latter part of
May to the early part of September, and it is probable that ovipo-
sition also takes place during this period, perhaps continuing a
little later. Mr. Garnett observed two sizes of larvae in the log.'
Only one pupal record was obtained, but judging from this,
and the condition of the insects on the various dates the log was cut
into or examined, it seems that the pupal period is about one
month, and the adult may remain in the log two weeks from the
time it forms until it eats its way out. Plate I, C — D, shows that
the beetle had to bore nearly an inch from the end of the gallery
in which pupation took place to the outside of the log.
In view of the relatively large numbers of such a rare beetle
collected by Mr. Garnett, Mr. Martin and the writer, Hubbard's
fears that the insect was about to become extinct are quite un-
founded. The two infested logs were found in the same canyon
but more than a mile apart.
Ent. News, Vol. XXIX, pp. 41-44, Feb. 1918.
\
Journal of Entomology and Zoology 67
HOLOTHUROIDEA
In sea-cucumbers the chief parts of the nervous system are
much as in other groups but the superficial and deep radial and
circum-oral systems are quite distinct from each other.
The more superficial system is composed of five strands in
an epineural cavity under the longitudinal radial muscles but well
in from the surface of the body. The oral ring circles the peris-
tome; at the base of the tentacles between its radial branches
there are strands, one for each tentacle ; other branches go to the
pharynx and intestinal tract. The epineural cavity seems not
present in some forms, possibly due to contraction of the animal.
The radial nerves end at the anal end of the body but there is no
special terminal tentacle. The radial nerves give off branches
to the tube-feet and also to the skin ; two nerve plexuses have been
recognized, a supei'ficial just under the epithelium and a deeper
one in the body-wall. Both of these networks receive some
branches from the radial nerve.
The deeper nerve ring or hyponeural divides into two strands
on the inside of each superficial radial nerve according to Hero-
uard, '87. This deeper system is chiefly motor while the super-
ficial system is sensory, a generalization which he extends to other
echinoderms. Branches from the deep system are said to supply
muscles of the body-wall and lantern region.
Among the earlier works dealing with the nervous system of
holothurians was that of Krohn, 1841, where the radial nerves
were noticed but little detail given. Semon, 1883, and especially
Hamann, show the general form and histological structure of the
nervous system. Herouard, '87-'89, brings out some points, espe-
cially emphasizing the motor and sensory divisions of the nervous
system, as already noted.
Gerould, '96, shows the nervous system in Caudina but little
is said about it. Clark in Synapta. 1898, shows the nervous system
in section. Red spots at the bases of the tentacles, the so-called
eyes, are figured.
Five radial nerves are recognized and smaller branches to the
tentacles. Each radial nerve is divided longitudinally into an
outer and inner band as in other forms, but unlike others has no
vessel of any kind accompanying the nerves and no spaces or
lacunae. Each tentacle nerve sends off branches to the digits so
that almost the whole surface of the tentacle becomes sensory. On
the bases of the tentacles and in the ectoderm over the body are
sense buds or tactile papillae such as described by Hamann, '83.
Under each of these is a small ganglion. From the lower side of
the circum-oral ring, between every two tentacles, a broad nerve
68
Journal of Kntomolog>' and Zoology
runs to the ectoderm of the oral disc and to the muscles of the
oesophagus.
Ackerman, 1902, gives figures of the nervous system in Cucu-
maria. Retzius, 1906, by means of the silver method gives a
mosaic picture of the epidermal cells. Between these cells are
small oval fields, the sense cells between the polygonal areas or
supportive cells. These are partly between two cells, partly be-
tween several supporting cells; they are not regularly arranged.
Reimers, 1912, discusses the development of Sintapta and gives
something of the nervous system. Haanen, 1914, in Mc.^othuria,
is not inclined to accept Herouard's (1890) suggestion that the
inner nerve band is chiefly a motor nerve. Very fine intestinal
nerves from the circum-oral nerve ring are found in this form as
well as the thicker nerves found by other observers. Every ten-
tacle and every foot has its own nerve, the first from the circum-
oral nerve ring, the second from the radial nerves. The foot
nerves are .029 inches broad and smaller and more circular in out-
line than the tentacle nerves. There seem to be at least some
Fig. 32. Nervous system of HoLOTHrROinEA. A. DiaRram of a sea-cu-
cumber showinfc superficial and deep central systems, branches to
tentacles and tube-feet and the inner and outer nerve plexus. B.
Section throujrh body-wall of Holothuria showing central band
in dark with nerve to a tube-foot. C. Nerve supply to tube-foot.
Hamann. D. Sense papilla of Synapta supplied by a nerve. Ha-
mann. E. Oral end of Synapta showing location of sense pores.
Pomona College, Clarcmont, California 69
motor and probably some sensory fibers in these. Sense cells and
an epithelial plexus were not clearly seen in this form. Retzius
found sense cells in the skin chiefly about the mouth opening, in
the tentacles and the tube feet. In this form the peripheral nerve
fibers were not found. Crozier. 1915, discusses the sensory reac-
tions of Holothuria surinamensls Ludwig.
The nervous system does not have to be intact for the act of
autotomy but it is more successfully carried out when it is unin-
jured.
The animals are reactive to tactile, vibratile, photic, and chem-
ical stimuli, and practically indifferent to heat in the way of a
sensation.
The parts of the body are sensitive in the following order,
beginning with the most sensitive: (1) tentacles, (2) anterior end,
(3) posterior end, (4) papillae, (5) pedicels (Podia), (6) mid-
body surface.
The tube-feet di.scs are positively stereotropic. This shows in
the righting reaction. The arms are photokinetic, negatively pho-
totropic ; they do not respond to increase in light intensity, but re-
spond negatively to decrease in light intensity. The whole surface
is sensitive in this way. The fluorei^cent skin pigment is possibly
concerned.
Dissolved substances representing those homologous to human
taste qualities for sour, bitter sweet, salt and alkaline, are effective
as stimuli.
Crinoidea
There are three distinct parts of the nervous system :
1. The superficial epidermal.
2. The deep oral system, according to the suggestions of
Delage and Herouard.
3. The deep aboral system.
The superficial oral system is much like the radial and circum-
oral system of starfish, with the nerve ring and radial nerves run-
ning down the surfaces of the ambulacral grooves in each arm with
branches to the surface and to the little elevations covered with
sense hairs.
The deep oral system according to Delage and Herouard's
interpretation is in the connective tissue under the epidermis and
consists of a central nerve ring and strands down each arm with
branches to the pinnacles.
The deep aboral system develops later than the oral in the
young form. It is in the center of the so-called chambered organ.
There is a central mass of nervous matter in the chamber; strands
run out from this towards the arms and fork but are united again,
70
Journal (if Entomnlo^y and Zonlony
to form a ring or pentagon of nervous tissue. From this ring
strands run out to each arm and branch and are distributed to the
arms, running embedded in the ossicles of the arms.
Carpenter, '66. and Marshall, '84, found that the aboral nerv-
ous system controls the movements of the animals. If the cham-
bered organ is destroyed the animal is paralyzed, but it will swim
readily or make the necessary movements just as well when the
whole ambulacral nerve ring and alimentary canal are removed.
Fig. 33. Nervous system of Crinoids. A. Diagram of a section through the
hotly of a crinoid showing nervous system hy heavier lines. B.
Diagram of a section of the nervous system of a crinoid. nerves in
black, after Marshall. C, D, and K. Diagrams of the central nerv-
ous system of Crinoids, after Marshall and Carpenter. F. Dia-
gram of the plan of the nervous system of a crinoid.
The commissural connectives between the aboral nerves co-
ordinate movements and if these are cut the arms move independ-
ently.
The position of the radial cords within the bony plates comes
about gradually from larval conditions when they are open,
trough-like grooves. These grooves gradually close in.
The cirri each have nerves from the central aboral nerve
mass. The arms, the cirri and the palps are tactile organs.
Hamann has shown nerve endings in the surface epithelium
as well as by means of little projections with fine hairs at their
ends.
Among the important contributions to the nervous .sy.stem of
this group are those of Carpenter, 1865-84, Teuscher, '76, Ludwig,
'77, Hamann, '87, Cuenot, '91. The pai)ers of Hamann, Carpenter,
Marshall and Haanen are among the most valuable contributions
to our knowledge of the nervous system.
Pomona Collefjc, Claremont, California 71
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Ackerman, A.
1900. Ueber die Anatomic unci zwittrigheit der Cucumaria laevigata.
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1872. Etudes generales sur le Systeme nerveux contrib. a I'hist. du
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Bronn, H. G.
1889. Tiereich. Bd. II. Abt. III. Echinodermen. 1-6 Holothuria; 7-16,
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1871-2. Ueber den Bau der Echinodermer. Sitz-Ber. Ges. Bef. ges.
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